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ANNALES CRYPTOGAMICI et PHYTOPATHOLOGICI 

Volume III 



BAZZANIA 



ANNALES CRYPTOGAMICI 
et PHYTOPATHOLOGICI 

[incorporating Annaks BryologiciJ 

edited by 

FRANS VERDOORN, PH.D. 

Managing Editor of Chronica Botanica. 'A New Scries of Plant Science 
Books', etc.; Bibliographer, Arnold Arboretum of Harvard University; 
Bot. Adviser, Board for the Neth. Indies; Hon. Sec., Bot. Section t 
Intern, Union of Biological Sciences; etc. 



Wij en konnen den Heer en maker van het geheel 
Al niet mccr vcrhecrlijken, als dat ivij in alle zaken, 
hoe klein die ook in onse blootc oogcn mogen sijn, als 
se maar levcn en ivasdom hebben ontfangen, sijn al 
wijsheit en volmaaktheit, met de uiterste vcrwondering 
sien uit stckcn. 

Antoni van Lecuivenlwek 



1946 

WALTHAM, MASS., U.S.A. 
Published by the Chronica Botanica Company 



The GENUS BAZZANIA 

in 
Central and South America 



by 



MARGARET FULFORD, PH.D. 

Assistant Professor of Botany, University of Cincinnati, 
Curator of He pa tics, Sullivan t Moss Society 




1946 

WALTHAM, MASS., U.S.A. 
Published by the Chronica Botanica Company 



First published 
By the Chronica Botanica Company 
of Waltham, Mass, U. S. A. 



rights reserved inchtd&ng the right to reproduce 
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PREFACE 



The study of Central and South American Hepaticae has been 
under way for about two hundred years and yet there is probably 
no other group of plants in that area in which the species are so lit- 
tle known or in which the taxonomy is in a worse state of confusion. 
This has not come about through any lack of interest in the plants 
in the field, for there is a long record of accumulated collections, but 
rather, it is due to the almost complete absence of critical revisions 
and monographs, without which an orderly taxonomy in any group 
is impossible. In addition to the usual accumulation of new species 
over this long period, there has also been an unfortunate rapid and 
voluminous production of "new species" by a few authors, among 
them STEPHANI, who described more than 1300 new species of he- 
paticae from tropical America alone. 

Monographic work is tedious and difficult because much of the 
original material and the obscure older literature is scarcely ob- 
tainable, yet the results reveal that the hepatics present as many 
and at least as varied problems of speciation, variability, distribu- 
tion, and ecology as are to be found in other groups. 

This work is humbly offered as the result of a taxonomic 
study of one of the larger American genera. The conclusions do 
not always agree with those of the earlier workers but it is sin- 
cerely hoped that those who use this volume will find the interpre- 
tations of the groups and species judicious and the text adequate. 



Annalea Cryptogomici et Phytopathologici, a new serial of 
which the present volume forms the third part, consists of 
memoirs (each forming a separate volume) devoted to gen- 
eral and systematic cryptogamy and to phytopathology. It 
continues Annales Bryologici. Vol. 1, GARRETT'S Root Dis- 
ease Fwngi was published in 1944, Vol. 2, HORSFALL'S Fungi- 
cides <md their Action in 1945. Vol. 4, CHESTER'S Cereal 
Rusts as Exemplified by the Leaf Ry^st of Wheat and Vol. 5, 
COPELAND'S Genera Filicum are in press. One or two vol- 
umes will be published every year at prices ranging from 
about $4.00 to $6.00. 

Annales Bryologici, a journal devoted to the study of 
mosses and hepatics, of which we published (in the beginning 
in cooperation with Messrs. Nijhoff) 12 volumes and 4 sup- 
plementary volumes, between 1927 and 1939 is now being con- 
tinued by the Annales Cryptogamici et Phytopathologici. 
Complete sets and single volumes of Annales Bryologici are 
still available at $4.00 a volume. 

The bryological exsiccati formerly issued by DR. FRANS 
VERDOORN: Bryophyta Arduennae Exsiccata (dec. 1-5, 
1927/29), Hepaticae Selectae et Criticae (11 series, 1930/39) 
and Musci Selecti et Critici (7 series, 1934/40) have all been 
sold out. 

The Chronica Botanica Co. also publishes "A New Series 
of Plant Science Books" and "CHRONICA BOTANICA, an inter- 
national Collection of Studies in the Method, Philosophy, and 
History of Biology and Agriculture". A catalogue with full 
information about these publications will be sent on request. 



ACKNOWLEDGMENTS 



The writer wishes to express her appreciation to all those persons and 
institutions who have assisted in any way with the accomplishment of this 
monograph. Plants from many herbaria have been studied, and the location 
of the individual specimens has been designated by the symbols enumerated 
as follows: (B) Botanical Garden and Museum, Berlin; (C) Field Museum, 
Chicago; (F) the herbarium of the writer; (G) herbarium of the University 
of Geneva (material from the STEPHANI Collection) ; (H) Cryptogamic Her- 
barium of Harvard University (material from the SCHIFFNEK, STEPHANI and 
TAYLOR Collections); (L) Rijksherbarium of Leiden; (NY) herbarium of the 
New York Botanical Garden (including material from the MITTEN Collection) ; 
(P) the private herbarium of the late F. M. PAGAN, Puerto Rico; (S) 
Botanical Garden, Goteberg, Sweden; (V) Natural History Museum, Vienna; 
(W) United States National Herbarium (including the collections of P. 
STANDLEY) ; (Y) herbarium of Yale University (including the private col- 
lection of A. W. EVANS). 

I also wish to acknowledge the generous cooperation of the Lloyd Library 
of Cincinnati in the loan of much valuable literature; the assistance and 
kindness of Dr. D. H. LINDER and associates of the Farlow Herbarium during 
visits there; and the generosity of the New York Botanical Garden in pro- 
viding facilities for research and several Research Scholarships at various 
times. Here Miss ROSALIE WEIKERT has been a valuable assistant in the care 
and handling of the plant material and Mrs. LAZELLA SCHWARTEN much aid 
in the Library. 

To Dr. A. W. EVANS of Yale University, for his kind cooperation, for 
his helpful suggestions and criticisms, and for reading much of the manuscript, 
the writer is deeply grateful. 

The author of this volume and the editor of the Annales Cryptogamici et 
Phytopatkologici would like to acknowledge their obligation to the Univer- 
sity of Cincinnati, which kindly awarded them a subsidy towards the cost of 
the illustrations. 



to 

ALEXANDER W. EVANS 



CONTENTS 



Introduction and History 1 

Characteristics of the genus Bazzania 5 

Description of the American species 7 

Key to the Subgenera 7 

Subgenus Bidentatae 8 

Key to the Species 8 

1. B. bidens (Nees) Trevis 9 

2. B. phyllobola Spruce 11 

3. B. Herminieri (Steph.) ..... 15 

4. B. cuneistipula (Gottsche & Lindenb.) Trevis. . 19 

5. B. platystipula Fulford 25 

6. B. roraimensis (Steph.) 27 

7. B. gracilis (Hampe & Gottsche) Stephani . . 30 

Subgenus Tridentatae 37 

Key to the Sections 37 

Section 1. Grandistipulae 38 

Key to the Species 38 

8. B. affinis (Lindenb. & Gottsche) Trevis. . 39 

9. B. pallide-virens (Steph.) . . . . 42 

10. B. stolonifera (Swartz) Trevis. . . 44 

11. B. chilensis (Steph.) 51 

12. B. taleana (Gottsche) .... 54 

13. B. denticulata (Lindenb. & Gottsche) 

Trevis 56 

14. B. quadricrenata (Gottsche) Pagan . . 60 

15. B. aurescens Spruce 63 

16. B. Glaziovii (Gottsche) .... 65 

17. B. Breuteliana (Lindenb. & Gottsche) 

Trevis 68 

18. B. acuminata (Lindenb. & Gottsche) 

Trevis 75 

19. B. diver sicuspis Spruce .... 77 

20. B. tricuspidata (Steph.) .... 79 

21. B. longistipula (Lindenb.) Trevis. . . 81 

22. B. latidens (Gottsche) .... 88 

23. B. longa (Nees) Trevis 90 

var. papillata (Steph.) . . 94 

24. B. jamaicensis (Lehm. & Lindenb.) Trevis. 96 

25. B. Wrightii (Gottsche) . . . .100 



Bazzania xv Contents 

Section 2. Connatae 103 

Key to the Species 103 

26. B. Fendleri (Steph.) . . . .105 

27. B. Schwaneckiana (Hampe & Gottsche) 

Trevis 106 

28. B. pycnophylla (Taylor) Trevis. . . 109 

29. B. Eggersiana (Steph.) Pag&n . . .110 

30. B. armatistipula (Steph.) .... 114 

31. B. cubensis (Gottsche) Pagan . . .116 

32. B. peruviand (Lehm. & Lindenb.) Trevis. 119 

33. B. Skottsbergii (Steph.) . . . .122 

34. B. novae-zelandiae (Mitten) Besch. & 

Massal 123 

Section 3. Fissistipulae 124 

Section 4. Appendiculatae 125 

Key to the Species 125 

35. B. asperistipula (Steph.) .... 125 

36. B. falcata (Lindenb.) Trevis. . . .129 

37. B. Hookeri (Lindenb.) Trevis. , . .135 

38. B. robusta Spruce 140 

39. B. Liebmanniana (Lindenb. & Gottsche) 

Trevis 142 

40. B. teretiuscula (Lindenb. & Gottsche) 

Trevis 144 

41. B. acanthostipa Spruce .... 151 

42. B. canelensis (Steph.) .... 152 

Section 5. Vittatae 155 

Key to the Species 155 

43. B. Spruceana Stephani .... 155 

44. B. Tayloriana (Mitten) . . . .157 

45. B. convexa (Thunb.) Trevis. . . . 159 

46. B. Stephani (Jack) 162 

Excluded species 165 

List of species not available for this study ... . 165 

Distribution 166 

Literature 168 

Index 171 



INTRODUCTION AND HISTORY 

The genus Bazzania was first proposed by S. F. GRAY (1821, 
704 as Bazzanius), when he recorded B. trilobata, formerly a part 
of the old genus Jungermannia, from England. Ten years later 
DUMORTIER (1831, 68) proposed the name Pleuroschisma, and soon 
after this, NEES VON ESENBECK (1833a, 96) introduced the name 
Herpetium for Jungermannia trilobata and allies ; in the year 1845 
NEES VON ESENBECK, LINDENBERG and GOTTSCHE (1844-47, 214) 
adopted the name Mastigobryum for the group. They divided the 
genus into three sections : A. Folia integra, including species with 
entire leaves; B. Folia bifida, including species with two-toothed 
leaves; and C. Folia tridenticulata, including species with three- 
toothed leaves. LINDENBERG and GOTTSCHE (1851) followed this 
same classification in a monograph of the genus, which at that time 
included 68 species. Later, STEPHANI (1886, 244), in a synopsis 
of the genus, classified the species under eleven sections, Integri- 
folia, Bidentata, Inaequilatera, Connata, Vittata, Parvistipula, 
Serrulata, Appendiculata, Fissistipula, Cordistipula and Grandi- 
stipula. SCHIFFNER (1893,101) described the genus under Bazzania 
and transferred the sections which STEPHANI had made for Masti- 
gobryum to this genus. Fifteen years later STEPHANI published 
his monograph of the genus Mastigobryum (1908, 413) in which 
he rearranged his former classification within the genus, raising 
three of his earlier sections to subgeneric rank and adding a fourth 
under which he included seven of the remaining sections. The 
Section Connata was here divided and included as subdivisions of 
the Sections Grandistipula and Serrulata. His revised classification 
follows : 

A. Integrifolia, 

B. Bidentata 

C. Tridentata 

1 . Parvis tipula 

2. Grandistipula 

a connata 
ft libera 
S. Serrulata 

a connata 
P libera 

4. Appendiculata 

5. Fissistipula 

6 . Cordistipula 

7. Vittata 

D. Inaequilatera 

A study of the American species (and 122 type specimens have 
been examined), indicates that in so far as the American species 



M. Pulford 2 Bazzania 

are concerned, the groupings of species which STEPHANI made were 
sometimes incorrectly assembled, and what is more important, 
were, in the case of several of the groups, purely artificial segrega- 
tions based on characteristics so variable that one species might be 
included in several sections. For this reason it seems advisable to 
revise the classification to more nearly conform with the natural 
groups within the genus. One of STEPHANI'S subgenera and sev- 
eral of his sections have been discarded and one of his older sec- 
tions revived. The reasons for these changes together with the 
characteristics of the subgenera and sections are discussed in the 
following. 

Integrifolia a subgenus in which the species have leaves 
which are not three-toothed or two-toothed, although the margins 
may be entire, serrulate or even dentate. Neither of the American 
species included here by STEPHANI actually belongs here, for both 
of them possess some leaves with three teeth more or less well 
developed ; and in addition, one of the species, M. Schwaneckianum, 
has the underleaf connate with one leaf. There are no representa- 
tives of this subgenus in America, and it may be that an examina- 
tion of the foreign species of this group will show that they also, 
should be included in the sections of the Subgenus Tridentata. 

Bidentata a subgenus made up of species in which the leaves 
are bidentate. This is a natural group of species which can be 
easily distinguished from those of the other subgenera. The in- 
dividual species show a closer relationship to one another than to 
any of the species within the other groupings. 

Tridentata a subgenus made up of species in which the 
leaves are three-toothed (sometimes only faintly so). The vast 
majority of the American species belong to this group. STEPHANI'S 
classification includes the following seven sections : 

1. Parvistipula made up of species with small underleaves. The 
American species which he included here are, for the most part, 
small, depauperate plants of species included in the following 
section. Only one, M. tricuspidatum, is a distinct species. The 
section has no distinctive characteristics except size of under- 
leaves, and this character alone is untrustworthy as a basis for 
segregation of a section. 

2. Grandistipula made up of species having leaves without a 
vitta or conspicuous auricles or appendages, and the teeth with 
entire margins. Two subdivisions are included by STEPHANI, 
libera, in which the underleaves are free from the leaves, and 
connata, in which the underleaves are connate with the leaves 
to a greater or lesser degree. The members of the connata 
group show a closer relationship to the members of the next 
section than to the libera group, and have been transferred. 
The members of the libera group, form a natural, closely re- 
lated unit. 



Bazzania 3 Introduction 

3. SernUata a section characterized by having the leaf margins 
more or less serrulate or spinose in the apical part. This sec- 
tion likewise is made up of two groups, libera and connata. 
The serrulate character is present to some degree throughout 
the group, but it is very variable and it is not always so dis- 
tinctive as one might wish. All of the American species of this 
section also have the underleaves connate with one or both 
leaves ; and at least some of the species of the libera group are 
actually connate. It is my opinion that the species having the 
underleaves connate with one or both leaves form a closely re- 
lated developmental group, and should be recognized as such, as 
STEPHANI did in his earlier classification (1886, 245). Since 
the serrulate condition of the leaf margins seems usually to be 
associated with the connate condition of the underleaves, but 
since not all of the species with connate underleaves have con- 
spicuously serrulate leaf margins it seems wise to redefine the 
group and to discard the Section Serrulata in favor of the 
Section Connata in order to avoid confusion. 

This Section would then include all connate species in the 
Subgenus (and also M. Schtvaneckianum from the Subgenus 
Integrifolia) . The species of the libera group if the leaves 
and underleaves actually are free, a point which should be care- 
fully investigated for all of the species in question belong in 
other already established groups. 

4. Appendiculata made up of species in which the ventral bases 
of the leaves are auriculate or appendiculate. This is a very 
distinct and natural segregation in which the characteristic 
tendencies of the group are exhibited in varying degrees by the 
members. 

5. Fissistipula including species in which the leaves are not ser- 
rulate along the apical margin, and are without ventral append- 
ages. The underleaves are deeply incised or long-toothed. The 
group appears to be natural and distinct. It is not represented 
in the Americas. 

6. Cordistipula including species with more or less cordate 
underleaves. The group is wholly artificial, since for the most 
part, it is made up of robust plants of species also included under 
the Grandistipula section, or rather slender plants of species be- 
longing to the AppendicuJata section. The Section has been dis- 
carded and the species for the most part reduced to synonomy 
in other sections. 

7. VMata made up of species in which the leaves all have con- 
spicuous vittas. This seems to be a natural assemblage of closely 
related species easily distinguished from those of the other 
sections. 

Inaequilatera the species of this subgenus have already been 
transferred to the genus Acromastigum (EVANS, 1934). 



M. Fulford 4 Bazzania 

As here revised, the organization of the genus Bazzania, based 
on a study of the American species, includes the following sub- 
divisions : 

Bidentatae 
Tridentatae 

Grandis tipulae 

Connatae 

A ppendiculatae 

Fissistipulae 

Vittatae 

Of the 444 species recognized in STEPHANI'S Monograph (1908- 
09; 1924), 115 species are reported from the Americas. Since that 
time other species have been added. Of the 122 species and varieties 
studied, 71 were reduced to synonomy, and one was transferred to 
another genus ; 22 species and varieties were not available. 

The species of North America north of Mexico have already 
been treated in detail ( FULFORD, 1936), so are not again included 
in this paper, since the southern boundary of the United States 
appears to mark the geographic boundary of the species of temper- 
ate North America on the south, and the tropical and subtropical 
species on the north. 



CHARACTERS OF THE GENUS BAZZANIA 
Bazzania 

Bazzania S. F. Gray, Nat. Arr. Br. PL 1: 704. 1821 (as Bazzanius). 

Pleuroschisma Dumort., Syll. Jungerm. 68. 1831. 

Herpetium Nees, Nat. eur. Leberm. 1 : 96. 1833. 

Herpetium, sect. Mastigobryum Nees, Nat. eur. Leberm. 3: 43. 1838. 

Mastigobryum Nees, Lindenb. & Gottsche, Syn. Hep. 214. 1845. 

Plants in large tufts or depressed mats, bright green to olive- 
green, golden yellow, reddish or brown ; stems filiform to robust, 
the vegetative branches of two sorts, leafy and flagelliform; the 
leafy branches from the ventral half of lateral segments, very 
rarely ventral, forming apparent dichotomies, the angle more or 
less constant for a species, in most cases less than 90 degrees, the 
incomplete leaf undivided, acute; cells of the leafy axis showing 
little differentiation internally, mostly thick-walled, the cortical 
layer similar to those of the medulla but of shorter cells, some- 
times deeply pigmented with brown; the flagelliform branches in- 
tercalary, arising mostly singly in the axils of underleaves, long, 
filiform, often branched, their branches intercalary, both lateral 
and ventral: rhizoids colorless, arising from the leaves of the 
flagelliform branches, the lower portions of the female bracts, and 
in some cases from the underleaves : the leaves incubous, alternate 
(rarely opposite), obliquely inserted, the line of insertion straight 
to strongly curved in its upper half, distant to imbricated, plane to 
deflexed, unsymmetrical, ovate to lanceolate, the dorsal base often 
strongly convex to cordate, the ventral base frequently more or less 
auricled, the apex in most species truncate, two- to three-toothed 
(in some four-toothed), in a few undivided; leaf margins in most 
cases entire, sometimes serrate to spinose-dentate, sometimes ciliate 
or appendiculate at the ventral base; the leaf cells quadrate to 
hexagonal in outline and more or less uniform except at the base 
where they are larger, or differentiated, with 6 to 12 series through 
the center of the leaf much enlarged, often of a different content, 
forming a vitta ; the trigones very small to large and well developed, 
often coalescing; the cell lumina rounded, or angular-rounded to 
stellate ; the cuticle smooth to verruculose : the underleaves distant 
to imbricated, quadrate, elongate or ovate, sometimes cordate at the 
base, the line of attachment transverse, slightly oblique, or re- 
curved, sometimes connate with leaves on one or both sides, the 
apex truncate, rounded-entire to two- to four-toothed or lobed, or 
variously incised, the margins entire, crenulate, spinose-dentate 
to ciliate ; the cells usually similar to those of the apical region of 
the leaf, all alike or those of the margin hyaline : dioicous, the male 
and female branches intercalary, in the axils of the underleaves; 
the male branches few to several on a stem, catkin-like, the brac- 
teoles slightly smaller than the bracts, the bracts in five or more 
series, ovate, concave to subcomplicate-con volute ; the apices trun- 
cate, bilobed to bispinose, rarely denticulate to entire; antheridia 



M. Fulford 6 Bazzania 

solitary or in pairs: female branches solitary, few to several on a 
stem; the bracts and bracteoles scarcely differentiated, in four or 
more series, closely imbricated, the innermost series largest, orbi- 
cular-ovate to ovate-lanceolate, the apex at least somewhat lobed, 
often two-, three-, or four-divided, the margins crenulate to 
laciniate-ciliate, the cells all alike or differentiated : the perianth to 
6 mm. long, ovoid-cylindrical, terete below, becoming three-keeled 
in the upper part, contracted at the mouth, of one layer of cells 
above, one to several layers below, the mouth of three ciliate to 
dentate lobes, usually contracted : the capsule oblong-ovoid, the 
wall usually of five layers of cells, the outermost layer with brown 
thickenings appearing as knots along the vertical walls, the inner- 
most layer with brown thickenings arranged as half-rings or bars 
on the inner tangential walls; the capsule stalk with a cortical 
layer of 16 large cells and a medulla of many smaller cells; the 
spores small, brown; elaters long, slender, bispiral; vegetative 
reproduction by means of vegetative shoots from the cells of or- 
dinary, as well as caducous leaves and underleaves. 

The plants of the West Indies, Mexico, Central America and 
South America give evidence of the same wide range of variation 
within a species as was pointed out (FULFORD, 1936) for the species 
of the United States and Canada. 

As in the case of the latter, these modifications seem to be of 
two sorts : the simpler, and by far the more common type, include 
those "temporary modifications" which have to do with the adapta- 
tions which occur in the individual plant in response to the condi- 
tions of the environment, and have been enumerated and discussed 
by BUCK (1928; 1929) in his studies of Scapania and Lophozia; 
the other sort seem to be, in part at least, hereditary and perhaps 
might be referred to as the various biotypic or genotypic expres- 
sions of the comprehensive Linnaean species. These "temporary 
modifications" include such characteristics as the variation in the 
amount of thickening in the secondary membranes, the degree of 
pigmentation of the cell membrane, the length of the internodes 
on the stem, and the size of the underleaves. Without doubt all of 
the characters listed above are influenced by the heredity of the 
species in question, but within limits, they may be strongly 
modified by environmental factors, particularly by the substrate, 
moisture, humidity, and the intensity of light. 

The writer has attempted to classify the species of Bazzania 
from the Americas with these concepts in mind. In many instances 
the results do not agree with those of earlier writers, particularly 
those of STEPHANI, but a study of all of the type material available 
has indicated that there are several instances in which he over- 
looked some of the most outstanding characteristics of the plant 
when making his classification. No doubt other workers will like- 
wise find inconsistencies and errors of judgment incorporated here. 
However, we like to agree with SPRUCE (1890, 116) when he states 
that, "in a genus where the numerous species are separated by such 
minute characters, two equally conscientious observers will often 
differ as to which are species and which are varieties". 



DESCRIPTION OF THE SPECIES 

The arrangement of the species follows in a general way that 
of STEPHANI in the Species Hepaticarum (1908-09; 1924) and the 
unpublished Icones, and in part both his subgeneric and sectional 
names, with feminine endings have been used. As has already been 
noted, p. 2, in some instances the characteristics of a species 
were not the same as the characteristics of the subdivision under 
which STEPHANI placed it, so that a transfer to a more suitable 
group has been necessary; other changes were necessary because 
certain of the sections, the Parvistipula and Cordistipula were 
purely artificial segregations based on extremely variable char- 
acters ; while still other changes have resulted from an attempt to 
bring together the members of a closely related unit. 

All of the American species of the genus Bazzania belong to 
two subgenera*, the Bidentatae and the Tridentatae. The latter 
group is further subdivided into the sections Grandistipulae, 
Connatae, Appendiculatae, and Vittatae. 

Key to the Subgenera 

Leaves predominantly two-toothed Bidentatae (p. 8) 

Leaves three-toothed, occasionally only faintly so ... Tridentatae (p. 37) 



"The American species which STEPHANI included in his Submenus Integri- 
folia actually belong to the Submenus Tridentatae, see p. 2, M. diversicuspe 
is a member of the section Grandistipulae, p. 38, and M. Schwaneckianum a 
member of the Connatae, p. 103. 



SUBGENUS BIDENTATAE 

Leaves predominantly bidentate, rarely tridentate or acute; 
underleaves attached in a straight line, not cordate at the base. 

The species form a well defined subgenus, all of them possessing 
the characters listed above. The American members are apparent- 
ly restricted to the tropics and subtropics, for there are no records 
for any of the species in the temperate zones. The leaves of B. 
ambigua of Canada and the northern United States are often two- 
toothed but most of the plants have some three-toothed leaves, for 
the species is definitely a member of the subgenus Tridentatae, 
closely allied to B. tricrenata and B. denudata. Of the seven species 
found in tropical America, B. bidens, B. cuneistipula, and B. 
gracilis are widespread in distribution, while the others seem to be 
restricted locally: B. phyllobola to northern South America, B. 
Herminieri to Guadeloupe, B. platystipula to Jamaica and Puerto 
Rico, and B. roraimensis to British Guiana. 

Key to the Species 

I. Stems coarse thread-like to delicate filiform, slender. 

2. Plants forming mats of very slender, dark brown, filiform stems, often 
without leaves. 

3. Leaves vittate, caducous 7. B. gracilis (p. 30) 

3. Leaves without a vitta, persistent (rarely absent), often reduced 
and inconspicuous, forms of . . . . 2. B. phyllobola (p. 11) 
2. Plants forming mats of coarser, green to brown stems; at least some 
leaves usually present, deflexed. 
3. Leaves in part caducous. 

4. Leaves with a vitta, the cell walls thin, the trigones small; 

underleaves small, distant 7. B. gracilis (p. 30) 

4. Leaves without a well differentiated vitta, the cell walls thin, 
the trigones very large, with convex sides; underleaves large, 
approximate to imbricated . . . 5. B. platystipula (p. 25) 
3. Leaves persistent. 

4. Underleaves large, approximate to imbricated; cells of the 
leaves with thin- walls and large, rounded trigones, 

5. B. platystipula (p. 25) 
4. Underleaves small, distant. 

5. Underleaves orbicular, entire or nearly so; trigones of the 

leaves large, rounded . . . 6. B. roraimensis (p. 27) 

5. Underleaves trapezoidal to subquadrate, two- to four- 

lobed at the apex 4. B. cuneistipula (p. 19) 

1. Stems not filiform, larger, the leaves spreading. 

2. Plants with leaves narrow-elongate to elongate-ovate. 

3. Margins of the teeth entire; underleaves subquadrate, the apical 
margins often faintly two- to four-lobed . . 1. B. bidens (p. 9) 
3. Margins of the teeth usually crenulate; underleaves with the 
margins variously and deeply incised, toothed, and lobed, 

2. B. phyllobola (p. 11) 



Bazzania 9 Bidentatae 

2. Plants with leaves unsymmetrically ovate. 

3. Leaves with a vitta of larger, thin-walled cells with conspicuous 
trigones, the teeth large. 
4. Teeth equal, the cells thin-walled, trigones small, 

7. B. gracilis (p. 30) 
4. Teeth unequal, the acroscopic tooth larger, the cell walls thin, 

the trigones large 3. B. Herminieri (p. 15) 

3. Leaf cells not differentiated, the teeth short, subequal; under- 
leaves trapezoidal to subquadrate, usually lobed at the apex, 

4. B. cuneistipula (p. 19) 

1. Bazzania bidens (Nees) Trevis. Mem. 1st. Lomb. 13: 415. 1877. 
Jungermannia tridens? Montagne, Ann. Sci. Nat. Bot. II. 3: 216. 1835, not 

Nees. 
Herpetium stoloniferum var. bidens Nees in Montagne, Ann. Sci. Nat. Bot. 

II. 14: 333. 1840. 
Mastigobryum bidens Gottsche & Lindenberg, in G. L. & N., Syn. Hep. 228. 

1845. 

Plants in mats, depressed to ascending, mostly large, yellow- 
green to brownish, becoming deeply pigmented with brown in the 
older portions : stems slender to robust, to 5 cm. or more in length, 
with leaves to 3.5 mm. broad; in longitudinal section the cells 
elongate, the medullary to 0.17 mm., the cortical shorter, both 
averaging 24/x in diameter, the vertical walls uniformly thickened 
and containing frequent pits, the end walls thin ; lateral branches 
frequent, mostly 5 mm. apart, diverging at a wide angle; flagelli- 
form branches numerous, long; rhizoids colorless, from the bases 
of the leaves of the flagelliform branches : the leaf insertion curved 
in its upper part; leaves distant to subimbricated, long, linear 
lanceolate, plane or only a little deflexed, 1.5 mm. - 2 mm. long, 
0.3 mm. - 0.5 mm. broad at the base, little narrowed to the trans- 
versely truncate, bidentate apex ; the dorsal margin convex from a 
straight base, the ventral margin straight to slightly concave, the 
base scarcely dilated, the apex two-toothed, the teeth very large, 
acute, mostly six to eight cells long and three to six cells wide, 
often divergent, the sinus deep, acute, the margins entire ; leaf cells 
thin-walled, the trigones large, with bulging sides, mostly coalesced, 
the cell lumina angular-rounded, the cuticle faintly verruculose; 
cells of the apical portion 32/x X 32/A, of the interior larger, and of 
the basal portion 42/A - 64/x X 32/x, a vitta not differentiated : under- 
leaves small, distant, subquadrate, a little wider than the stem, 
attached in a straight line, 0.35 mm. - 0.40 mm. long X 0.31 mm. - 
0.35 mm. wide, the lateral margins nearly straight, entire, the apex 
undulate, variously toothed or lobed, the cells as in the apical por- 
tion of the leaf : leaves of the flagelliform branches scale-like, ovate, 
acute to bifid : male branches frequent, several on a stem ; the bracts 
concave, ovate, bifid, crenulate, the cells to 48/x long X 24/x wide, 
the bracteoles similar, smaller ; antheridia in pairs : female branches 
solitary, several on a stem (very poorly preserved) ; bracts of the 
intermediate series divided to one-third into usually three laciniae, 
the margins serrate to laciniate : sporophyte not seen. 

HABITAT : Over rocks, logs, and on trunks of living trees. 
The distinguishing characteristics of the species are its rather 
robust habit, the yellow-brown color, and the plane, long, linear- 




FIG. 1. Bazzania bidens (Nees) Trevis. 1. Portion of a stem, ventral 
view, X 12. 2. Underleaves, X 30. 3. A cell from the apical portion of a 
leaf, X 400. 4. Portion of a stem and leaves, dorsal view, X 40. 5. Leaves, 
X 25. 6. A tooth of a leaf, X 200. 7. Cells from the dorsal margin of a leaf 
near the base, X 200. 8. Cells of the basal portion of a leaf, X 200. 9. Under- 
leaf, X 60. 10. Portion of a transverse section of a stem, X 310. 11. Female 
bracts of an intermediate series, (immature) X 30. Nos. 1-3 drawn from the 
type material from French Guiana; 4-11 from material collected by EVANS 
in Puerto Rico. 



Bazzania 11 Bidentatae 

lanceolate, strongly bidentate leaves which are usually approximate. 
The plants are often quite brittle. (FIGURE 1, nos. 1-11). 

While there is considerable variation in the size of the plants 
and the size of the trigones of the leaves, the leaves are always 
narrow, elongate, and the trigones always large with convex sides 
(see FIG. 1, nos. 1, 3, 5 and 6). 

The female branches were poorly developed and in a poor state 
of preservation in all the examples available. The bracts of the 
intermediate series are broadly ovate and are to one-third divided 
into two or three laciniae (FiG. 1, no. 11). The lateral margins 
are serrate, dentate or laciniate. This agrees in general with 
STEPHANIES figure in the Icones, Mastigobryum no. 25. He shows 
the bracts of the innermost series as being two-thirds divided into 
two, dentate-ciliate laciniae. Unfortunately, bracts of this series 
were too poorly developed for comparison. 

SPRUCE (1885, 371) described and later distributed in the 
Hepaticae Spruceana, plants from Brasil as B. bidens, which are 
of an entirely different habit from those of the type collected by 
LEPRIEUR in French Guiana. SPRUCE'S plants represent a large 
form of B. gracilis and are discussed further under that species. 

DISTRIBUTION : Dominica: without locality, Elliott, cited by Stephani 
(1908, 440). Guadeloupe: without locality, PHerminier, cited by 
Stephani (1908, 440); without locality, Duss, cited by Stephani (1908, 440). 
Duss 1096 (NY) should be referred to B. Herminieri. Martinique: 
without locality, Husnot, PI. Antilles, 241b (H,G). Some of the packets also 
contain a little B. longistipula (NY). Puerto Rico: without locality, 
Schwanecke (G) ; without locality, Sintenis 37, cited by Stephani (1888a, 
279) ; El Yunque, Luquillo Mountains, Evans 51 (Y, NY) ; Sierra de Naguabo, 
Shafer 3750 (NY); El Yunque, Pagan 494 (NY). French Guiana: 
without locality, Leprieur, the type (G) ; without locality, Hb. Hooker (NY). 
Peru: without locality, Lechler (G). Guiana : without locality, 
Montagne (NY). 

The specimens from Brasil distributed by SPRUCE in Hepaticae Spruceana: 
Amaz. et And. as B. bidens are B. gracilis. 

REFERENCES: Lindenberg and Gottsche (1851, 87, pi. 15); Hampe and 
Gottsche (1852, 346) ; Husnot (1875, 3) ; Spruce (1885, 371) ; Stephani (1888a, 
279; 1908, 439; Icones, Mastigobryum no. 25) ; Bescherelle (1893, 186) ; Pagan 
(1939, 38). 

2. Bazzania phyllobola Spruce, Trans. & Proc. Bot. Soc. [Edin- 
burgh] 15: 372. 1885. 

Mastigobryum phyttobolum Stephani, Spec. Hep. 3: 439. 1908. 
Bazzania bidens var. dissodonta Spruce, op. cit. p. 371. 
Bazzania dissodonta Spruce ms. 

Mastigobryum dissodontum Stephani, Trans. Linnean Soc. Bot. II. 6: 98. 
1901-05. 

Plants slender, brownish green to deep reddish brown in the 
older portions: stems to 6 cm. long, with leaves to 3 mm. broad, 
depressed to ascending : stem cells in longitudinal section elongate, 
to 0.17 mm. long, the cortical shorter, both averaging 15^ in 
diameter, the vertical walls uniformly thickened and containing 
frequent pits, the end walls thin ; lateral branches frequent, 5 mm. 



M. Fulford 12 Bazzania 

or more apart, diverging at a wide angle; flagelliform branches 
frequent, long ; rhizoids colorless, scarce, on the bases of leaves of 
the flagelliform branches: leaf insertion little curved in its upper 
part; the leaves approximate to subimbricated on well developed 
plants, deflexed when dry, unsymmetrical, elongate-ovate, ascend- 
ent, 0.75 mm. - 1.5 mm. long, 0.5 mm. broad at the base, narrowed 
a little to the transversely truncate, bidentate apex; the dorsal 
margin arched from a scarcely rounded base, covering approxi- 
mately one-half the stem, the ventral margin nearly straight, the 
base scarcely dilated, the apex two-toothed, the teeth large, acute, 
six to eight cells long, five to seven cells broad, the sinus deep, 
acute, the margins crenulate, serrate or even shortly denticulate; 
leaf cells thin-walled, the cell lumina rounded, the trigones small, 
conspicuous, rarely coalesced, the cuticle verruculose; the cells 
of the apical region mostly 20/x X 20^, those of the base 
36/i - 40/x X 24/x, a vitta not differentiated : underleaves distant to 
approximate, more or less quadrate in outline, attached in a straight 
line, little broader than the stem, mostly 0.26 mm. - 0.32 mm. long 
and broad, the apex irregularly divided into irregular, long or 
short, blunt to acute teeth, the lateral margins entire, or variously 
toothed, the cells mostly as in the apical part of the leaf; leaves 
of the flagelliform branches scale-like, ovate, acute to shortly bifid : 
female branches frequent, solitary* one to several on a stem; the 
bracts and bracteoles similar, the outer series small, ovate, one- 
third divided into two blunt teeth, the lateral margins serrate ; the 
intermediate and innermost series (immature) ovate, to one-half 
or more divided into two or three long, slender, laciniae, one or 
two cells broad, twelve or more cells long, the lateral margins 
serrate to ciliate: perianth (immature) mouth densely long-ciliate : 
male bracts and sporophytes not seen. 

HABITAT : On tree bases, forests in the mountains. 

The distinguishing characteristics of the species are its green- 
ish brown to reddish stems, the deeply two-toothed leaves, the 
crenulate to denticulate margins of the teeth; or long, reddish 
stems with smaller, scattered, acute, two- or three-toothed leaves 
of various sizes ; the strongly and irregularly toothed underleaves 
(sometimes entire on filiform stems) ; and the female bracts 
divided in the upper part into long, slender, ciliate laciniae. (FlG. 
2, nos. 1-19). 



FIG. 2. Bazzania phyllobola Spruce. 1, 2, 3. Portions of stems, ventral 
view, X 30. 4. A cell from the apical portion of a leaf, X 400. 5. A lacinia 
from a female bract of the innermost series, X 100. 6. Cells of a lacinia of 
this series, X 310. 7. Cells from a portion of the mouth of the perianth, 
X 310. 8. Outermost female bract, X 30. 9. Portion of a stem, ventral view, 
X 30. 10. Dorsal side of stem to show leaf attachment, X 30. 11. A cell from 
the apical region of a leaf, X 400. 12. Apical tooth of a leaf, X 310. 13. Cells 
from the dorsal margin of a leaf, X 310. 14. Cells from the base of a leaf, 
X 310. 15. Cells from the apical portion of an underleaf, X 310. 16. Female 
bracts of an intermediate series, X 30. 17. Female bracts (munature) of the 
innermost series, X 30. 18. A lacinia from a bract of this series, X 100. 
19. Portion of the perianth mouth (immature), X 100. Nps. 1-8 drawn from 
a portion of the type material; 9-19 drawn from the material of B. dissodonta 
Spruce ms. collected by SPRUCE on Mt, Guayrapurina. 







Ou&c 



M. Fulford 14 Bazzania 

The original material of B. phyllobola distributed by SPRUCE is 
made up mostly of very long, red brown stems with distant leaves 
(FiG. 2, nos. 1-4), some of which may have been caducous, because 
stems without leaves except at the tips occasionally occur. The 
leaves are mostly linear, shortly bilobed or acute, and usually small. 
On a few of the stems occasional leaves are well developed and 
identical with those of B. bidens var. dissodonta in shape, cell 
structure, and configuration of the apical margin (compare FIG. 
2, nos. 1 and 9). The underleaves, for the most part, are very 
small and entire or faintly toothed or lobed on the apical margin, 
but all gradations up to the large, toothed sort, characteristic of 
B. bidens var. dissodonta can be found with little difficulty on stems 
where the leaves are more or less well developed. Flagelliform 
branches are produced in abundance and very often become leafy. 
Jn addition, the main stems with well developed leaves may become 
smaller, the leaves lose their characteristic shape and become 
linear-bifid or acute, and the underleaves become entire or nearly 
so, so that the stems assume a flagelliform appearance. Many of 
the plants, even those with few leaves, had produced several female 
branches. The archegonia were unfertilized and the bracts had 
not matured. The intermediate and innermost bracts are ovate, 
divided into long, slender, ciliate laciniae, and are identical with 
those of B. bidens var. dissodonta (compare FIG. 2, nos. 5-8 with 
nos. 16-19). They are in agreement with those of STEPHANIES 
1 COUCH, Mastigobt^yum nos. 28 and 33. According to the labels, 
SPRUCE collected both his B. phyllobola and B. bidens var. dissodonta 
in the Peruvian Andes on Mt. Guayrapurina. 

B. bidens var. dissodonta (FiG. 2, nos. 9-19) represents the 
more typical, normally developed aspect of the species. Here the 
plants are large, conspicuously leafy, greenish brown instead of 
rod-brown, and only rarely do leafy stems become fiagelliform in 
appearance. The leaves are unsymmetrically ovate, well developed, 
and deeply two-toothed. The teeth are large, more or less spread- 
ing, and the margins are characteristically crenulate or even denti- 
culate. Three-toothed leaves are occasionally present. The leaf 
cells do not differ from those of the leaves of B. phyllobola. The 
enlarged cells of the interior suggest a vitta, although they are not 
particularly conspicuous. 

The underleaves are very characteristic. They are broader than 
the stem, irregularly and deeply toothed and lobed, and tend to be 
quadrate in outline. 

The female branches, which are numerous, possess bracts (im- 
mature), which are ovate in outline and to one-half divided into 
long, ciliate laciniae. A few of the plants in the mat are smaller, 
with more distant leaves and are more nearly like the original 
material of B. phyllobola. The perianth mouth is long ciliate. 

The species, when well developed, is similar in many ways to 
H. bklcHs but can readily be distinguished from it because of the 
crenulate to denticulate margins of the teeth of the leaves, and the 



Bazzania 15 Bidentatae 

variously incised and toothed underleaves. The female bracts of 
the two species are very similar. The small forms of the species are 
similar to B. gracilis but can usually be distinguished by the ab- 
sence of a well defined vitta in the small leaves. 

DISTRIBUTION : British Guiana: without locality, McConnell & 
Quelch as M. dissodontiun, cited by Stephani (1901-05, 98). Brasil : 
without locality, Ule (H) ; Santas, Horeau, cited by Stephani (1908, 439). 
Colombia: cited by Stephani (1901-05, 98). Peru: Mt. Campana, 
Spruce as B. bidens var. dissodonta, cited by Spruce (1885, 371); Mt. Guay- 
rapurina, Spruce as B. dissodanta, Hepat. Sprue. (NY); Mt. Guayrapurina, 
Spruce, the type, Hepat. Sprue. (NY). 

The plants collected by STANDLEY 48091 in Costa Rica (Herzog 1938, 19) 
are B. gracilis. 

REFERENCES: Herzog (1938, 19) ; Stephani (1886, 134; 1901-05, 98; /cones, 
Mastigobryum nos. 28, 33). 

3. Bazzania Herminieri (Steph.) comb. nov. 

Mastigobryum Herminieri Gottsche in HUSNOT, Rev. Bryol. 2: 3. 1875. (nomen 
nudum) ; in STEPHANI, Hedwigia 25: 8. pi. 4. fig. 3-6. 1886. 

Plants growing in depressed to ascending mats, mostly large, 
dark brown in the older portions, light yellow-green to brown at 
the growing tips : stems slender to robust, to 10 cm. or more long, 
with leaves 2 mm. - 2.5 mm. wide, mostly ascending to erect; in 
longitudinal section the cells elongate, the medullary cells averag- 
ing 0.18 mm. long, the vertical walls uniformly thickened and con- 
taining frequent pits, the end walls thin, the cortical cells shorter, 
the walls thicker, deeply pigmented with brown, both averaging 
18/i in diameter: lateral branches infrequent, diverging at a wide 
angle; flagelliform branches frequent, long; the line of leaf- 
insertion curved in its upper half; the leaves imbricated, plane to 
convex, often becoming strongly deflexed on drying, so that dried 
stems appear to be laterally compressed, unsymmetrically ovate, 
more or less falcate, to 1 mm. long, 0.6 mm. - .72 mm. wide at the 
base, narrowing to 0.3 mm. at the bidentate apex : the dorsal margin 
strongly convex from a curved base, the ventral margin straight 
to concave, the apex two-toothed, the acroscopic tooth the larger, 
acute, mostly eight to ten cells broad at the base, six to twelve cells 
long, widely spreading, the sinuses lunulate to deep and acute, the 
margins entire, somewhat undulate; leaf -cells of two sorts, those 
of the marginal and apical portions more or less isodiametric, 
averaging 18/*, the trigones large with bulging sides, often becom- 
ing coalesced, the cell cavity angular-rounded ; those of the median 
and basal portions of the leaf elongate, forming a distinct vitta 
near the base, averaging 50/x long X 25/x wide : the cuticle verru- 
culose: underleaves attached in a straight line, distant, becoming 
subimbricated near the growing point, subquadrate, as broad or 
broader than the stem, 0.32 mm. - 0.4 mm. wide, 0.24 mm. - 0.54 
mm. long, the apex faintly to deeply two- to four-lobed, the lobes 
rounded-entire, the sinuses mostly narrow-acute, the lateral margins 
straight to convex, undulate: leaves of the flagelliform branches 
scale-like, convex, to 0.18 mm. long, ovate, the apex obtuse to biden- 
tate : male branches usually several on a stem ; the bracteoles small, 
rectangular in outline, somewhat convex from below, 0.4 mm. - 



M. Fulford 16 Bazzania 

0.45 mm. long X 0.24 mm. - 0.3 mm. wide, the lateral margins 
straight, entire, the apex faintly to strongly two- to four-lobed, 
with shallow, acute sinuses; the bracts larger, broadly quadrate- 
orbicular, strongly convex from below, averaging 0.56 mm. long, 
to 0.7 mm. wide, the lateral margins bulging, entire to crenulate, 
the apex with two to four short teeth and lunulate to acute sinuses ; 
the cells of the apical portion with large, confluent trigones, the 
cell-lumina stellate, those near the base rectangular, thick-walled, 
the trigones not conspicuous; antheridia usually in pairs: female 
branches frequent, solitary, one to several on a stem; the outer- 
most bracts averaging 0.48 mm. long X 0.32 mm. broad, the 
margins entire, the apex mostly two-lobed, the cells elongate except 
at the margin, thick-walled ; the intermediate bracts larger, mostly 
0.72 mm. long X 0.5 mm. wide, the upper third divided into three 
or four narrow, mostly blunt, entire laciniae, the lateral margins 
entire to obscurely dentate, the cells elongate, 30/A X 12/A in the 
upper portion, 45/x X 18/x near the base, thick-walled, often ir- 
regularly so, the trigones large: perianth to 6 mm. long, ovoid- 
cylindrical, contracted at the mouth, of one layer of cells throughout, 
at the mouth divided into numerous, unequal, mostly narrow, short 
to long-pointed laciniae, the margins entire to serrate, the cells 
20/x - 36/x long, mostly 15/i wide, the walls thick, cells of the basal 
portion mostly 65/x, X 18/x, the walls unevenly thickened: sporo- 
phyte not seen. 

HABITAT: "Abundant on rocks", HUSNOT, PL de Antilles; on 
rocks, tree bases and stones. 

The distinguishing characteristics of the plant are its medium 
size, the elongate, vittate leaves which tend to be slightly falcate 
and which are mostly unequally two-toothed at the apex; the thin 
cell walls and conspicuous, often confluent trigones; and the 
quadrate mostly four-lobed underleaves. (FiG. 3, nos. 1-20). 

The leaf apex is quite variable. It is usually unequally two- 
toothed, with the acroscopic tooth the larger, but in some of the 
leaves the teeth tend to become equal or to disappear completely. 

The vitta is well defined. It consists of five or six rows of cells 
running through the middle of the leaf from the base half-way to 
the apex. It is more or less sharply distinct from the cells of the 
ventral margin because of the difference in the size of the two, but 
the distinction between it and the cells of the dorsal margin is less 



FIG. 3. Bazzania Herminieri (Steph.) Fulford. 1. Portion of stem, 
dorsal view, X 30. 2, Portion of stem, ventral view, X 12. 3. Leaf, X 30. 
4. Underleaves, X 30. 5. Cells from the apical portion of an underleaf, 
X 260. 6. Apical tooth of leaf, X 260. 7. Cells from dorsal margin of leaf, 
X 260. 8. Cells from the ventral margin of leaf, X 260. 9. Cells from the 
base (vitta) of a leaf, X 260. 10. Portion of a longitudinal section of a 
stem, X 260. 11. Portion of a cross-section of stem, X 260. 12. Female 
bract of the outermost series, X 30. 13. Female bracts of intermediate series, 
X 30. 14. Female bract of the innermost series, X 100. 15. Apical tooth of 
bract, X 100. 16. Portion of mouth of perianth, X 30. 17. Portion of mouth 
of perianth, X 100. 18. Cell from near the base of perianth, X 260. 19. Male 
bracteole, X 30. 20. Male bracts, X 30. Nos. 1-9, 11-13, 19, and 20 drawn 
from material of Husnot, PL Antilles, no. 211; nos. 10, 14-18 drawn from 
material collected by THEEMINIEB, in the Herbarium of GOTTSCHE. 



M. Fulford 18 Bazzania 

pronounced because of a more gradual transition (FiG. 3, nos. 2, 
3,7-9). 

The underleaves are quadrate, a little broader than the stem, 
and lobed along both the lateral and apical margins in well devel- 
oped plants (FiG. 3, nos. 2 and 4). Many variations in size and 
form occur. In some, the lateral margins tend to become straight 
and entire, and the lobing of the apices less distinct. Quite often 
in some of the less well developed forms the underleaves are much 
broader at the apex than at the base and cannot readily be 
distinguished from those of B. cuneistipula. 

The thickness of the walls of the stem cells varies with the age 
of the plant, as does the degree of brown pigmentation. The size 
of the trigones and the thickness of the cell-walls in both leaves 
and bracts also show a direct correlation with age and exposure. 
The deposit is often very irregular on the inner walls of the cells 
of the female bracts and the perianth (FiG. 3, no. 18). 

Plants collected by Duss, no. 1096, among mosses and 
Sphagnum, in Guadeloupe, vary considerably in appearance from 
the usual forms. They grew erect in compact tufts. For the most 
part, the stems are more slender than in plants from other 
habitats, and the leaves, while well developed, are distant or only 
approximate. 

GOTTSCHE designated several forms in the material from 
Guadeloupe. The forma latior G. with shorter leaves is less well 
developed than the type, and has the vitta not so conspicuous as in 
the larger plants. The forma applanata G. is of the usual size, 
with plane, spreading leaves which are unequally bidentate and 
have conspicuous vittae. The variety brevifolia G. is a depauperate 
stem with the leaves very distant. All of them could no doubt be 
duplicated in any large collection of the species. 

The plants are somewhat similar to those of B. bidens and B. 
phyllobola. The large, plane, elongate leaves with large equal teeth 
of B. bidens, and the large crenulate teeth and the conspicuously 
incised and toothed underleaves of B. phyllobola will readily 
identify those species. The female bracts of the three species are 
also distinctive. 

DISTRIBUTION : Guadeloupe: Souf riere, I'Herminier, distributed in 
Husnot, PL Antilles, 211, the type (H, Y, NY) ; plants of the same collection 
from the Herbarium of GOTTSCHE 23, 36 (NY) ; plants of the same collection 
as var. brevifolia G., forma applanata G., and forma latior G. (H) ; Souf riere, 
Duss 1034, 1077 and 1096 as M. tenerum (NY), nos. 62, 98, 102, 122, 302, 309, 
310, 321, 328 (H). Martinique: Mt. Pelee, Duss 348 as M. varia- 
bile (NY). 

REFERENCES: Bescherelle (1893, 186); Stephani (1903, 22; 1909, 437; 
Icones, Mastigobryum no. 31). 



Bazzania 19 Bidentatac 



4. Bazzania cuneistipula (Gottsche & Lindenb.) Trevis. Mem. 1st. 

Lomb. 13: 414. 1877. 

Mastigobryum cuneistipulum Gottsche & Lindenberg, in G. L. & N., Syn. Hep. 

225. 1845. 

Mastigobryum tenerum Gottsche & Lindenberg, loc. cit. 
Bazzania tenera Trevis. loc. cit. 

Mastigobryum variabile Hampe & Gottsche, Linnaea 25: 348. 1852. 
Mastigobryum brevifolium Gottsche, Ann. Sci. Nat. Bot. V. 1: 141. 1864. 
Mastigobryum corticola Stephani, Spec. Hep. 3 : 467. 1908. 

Plants scattered or growing in depressed mats, small, dull gray- 
ish green to yellow-brown: stems slender, to 5 cm. or more in 
length, with leaves mostly 1 mm. - 1.5 mm. wide, prostrate to 
scarcely ascending; in cross-section the cells averaging 18//. in 
diameter, the walls thickened, the pits numerous; lateral branches 
frequent, 3 mm. or more apart, diverging at a wide angle; ventral 
branches abundant, long, often branched, flagelliform, sometimes 
becoming leafy; rhizoids colorless, abundant on the leaves of the 
flagelliform branches: the line of leaf insertion a little curved in 
its upper half; the leaves distant to subimbricated, ascending, plane 
to more or less convex, becoming defiexed or wrapped around the 
stem when dry, unsymmetrically ovate, mostly 0.7 mm. - 1 mm. 
long, 0.4 mm. - 0.5 mm. wide at the base, narrowing to the biden- 
tate apex; the dorsal margin convex from a straight or rounded 
base, extending over one-third of the stem, the ventral margin 
nearly straight; the apex normally two-toothed, the teeth mostly 
subequal, two to five cells long, spreading, the sinus broad, lunulate 
to acute; the leaf cells thin-walled, the trigones conspicuous, often 
confluent, the cell lumina angular-rounded, the cuticle verruculose ; 
cells of the apical portion and dorsal base 18/* - 27/t in diameter, 
those of the median portion larger, those of the base to 36/x X 20/i, 
a vitta not differentiated: underleaves distant, attached in a 
straight line, squarrose, as broad or broader than the stem, sub- 
quadrate, truncate-trapezoidal to quadrate-orbicular, 0.24 mm. - 
0.36 mm. long and wide, often narrowing a little to the base, the 
lateral margins straight to slightly undulate, the apex mostly four- 
lobed, the lobes broad, rounded, the sinuses shallow, narrow, acute 
to rounded, the cells as in the apical portion of the leaf: leaves of 
the flagelliform branches scale-like, appressed, ovate, mostly 0.12 
mm. long, the apex rounded, acute to bidentate: female branches 
frequent, one to several on a stem, the bracts and bracteoles similar ; 
the outermost series small, rectangular in outline, usually shortly 
two-lobed, the margins entire; the intermediate series ovate, 
averaging 0.64 mm. - 72 mm. long, 0.4 mm. broad at the base, 
approximately one-fourth divided into usually two, sometimes 
three, narrow, entire, blunt to acute teeth, the lateral margins 
crenulate, often undulate, the cells elongate, those of the apical 
portion averaging 25/A long X 15/x wide, of the basal portion 54/A 
X 18/x,, the walls mostly uniformly thickened, trigones not con- 
spicuous; the innermost series (not mature) ovate, to 0.96 mm. 
long, the upper fourth divided into two or three, mostly entire 
laeimae, the lateral margins cromilato to dentate, the cells elongate, 
more or less uniform throughout, in the median portion averaging 
72/x X 18/A, the walls rather thin, the trigones indistinct: the 



M. Fulford 20 Bazzania 

perianth (immature) mouth divided into numerous laciniae, often 
ciliate: male branches and sporophytes not seen. 

HABITAT : On trees and logs in woods. 

The distinguishing characteristics of the plant are its small 
size, the shortly bidentate leaves with mostly lunulate sinuses and 
large thin-walled cells with conspicuous trigones; and the trape- 
zoidal to round-quadrate underleaves with mostly four-lobed apices. 
(FiGS. 4, nos. 1-18; 5, nos. 1-14). 

While most of the underleaves of the type material show the 
straight lateral margins and the Cuneate tendency as shown in 
FIG. 4, no. 9, others show no decrease in width from the apex to 
the base (FiG. 4, no. 2), and in some instances the margins are 
convex or bulging and to a slight degree undulate. The under- 
leaves on the plants examined, in no case exhibited so marked a 
difference in width between apex and base, as is indicated by the 
figures in the Species Hepaticarum (LiNDENBERG and GOTTSCHE, 
1851, pi. 13). The apices instead of being usually more or less 
deeply bifid, as they have indicated, are only slightly depressed, or 
more often, nearly straight or inconspicuously four-lobed. The 
same variations are even more pronounced on the less well developed 
parts of a plant. 

The degree of brown pigmentation and the thickness of the cell 
walls vary with the age and habit of the plant. This is noticeable 
in both the leaves and in the stem. Stem sections from near the 
growing tip showed only one layer (cortical) of cells with ex- 
tremely thick, light brown walls, and a medulla of many thin-walled 
cells, while sections from much older parts of the stem showed a 
much wider border of thick-walled cells strongly pigmented, and 
only a few thin-walled cells. In older leaves the cuticle is strongly 
verruculose. Those plants from the mountains are usually brown 
except at the growing tips. 

The leaves are prevailingly two-toothed at the apices, but very 
often show modifications as indicated in FIG. 4, no. 4. There is 
much variation even in the two-toothed condition. In some, the 
apex is narrow and the sinus narrow and deep, in others the apex 
is broad, the teeth are broad and short, and the sinus is lunulate. 
The teeth are long or short, rounded to acute, and equal to strik- 
ingly unequal. The leaves are mostly broader and shorter than 
LINDENBERG and GOTTSCHE have indicated (1851, pi. 13, fig. 4). 

FIG. 4. Bazzania cuneistipula (Gottsche & Lindenb.) Trevis. 1. Por- 
tion of a stem, dorsal view, X 30. 2. Portion of a stem, ventral view, X 30. 
3. Leaf, X 30. 4. Variations in leaf apices, X 30. 5. Cells from the apical 
portion of a leaf, X 260. 6. Cells from the dorsal margin of a leaf, X 260. 
7. Cells from the ventral margin of a leaf, X 260. 8. Cells from the basal 
portion of a leaf, X 260. 9. Underleaves, X 30. 10. Cells of the upper part 
of an underleaf, X 260. 11. Portion of a cross-section of a stem, X 260. 
12. Female bract of the outer series, X 30. 13. Female bracts of an intermedi- 
ate series, X 30. 14. Upper part of a female bract of the intermediate series, 
X 260. 15. Female bracts of the innermost series, X 30. 16. Outline of a 
portion of the mouth of a perianth, X 30. 17. Portion of perianth mouth. 
X 260. 18. A cell from the base of a bract showing irregular thickenings of 
the wall, X 260. Nos. 1-18 drawn from a portion of the type material. 




^nSW 




M. Fulford 22 Bazzania 

Most of the cells of the bracts show a uniformity in the thick- 
ness of the cell walls, but often (in older bracts), the walls of many 
of the basal cells show wart-like lumps of thickening irregularly 
scattered over the inner surfaces (FiG. 4, no. 18). The number, 
size, and shape of these thickenings vary in different cells. The 
same condition has been observed in the cells of the female bracts 
and perianths, and even the stems of many of the species of the 
West Indies. EVANS (1933, 74) has described the same condition 
in the cells of the perianth of B. loricata from Sumatra. 

The species is quite similar in many ways to the more northern 
B. ambigiut of the northwestern United States and Canada, which 
often has many bidentate leaves, and underleaves with mostly 
straight sides. The two species are approximately the same size 
and the same color, and have the same prostrate to ascending habit 
of growth. However, the leaves of B. cuneistipula are larger, and 
are composed of larger cells. They also have a greater number of 
enlarged cells in the basal portion. The two are further distin- 
guished by differences in the female bracts and in the perianths. 

Some of the plants from Jamaica show certain characteristics 
which do not occur in plants of the type collection. They are mostly 
yellow-brown, smaller, and have a coarse filiform appearance. The 
leaves are strongly deflexed, much more so than in the type, and 
curve around the stem when dry. The underleaves vary from 
trapezoidal to quadrate-orbicular. 

Few comments concerning the size and habit of M . tenerum can 
be made from the type 1 since so little material has been available 
for study. Most of the other specimens also are either very meager 
or in a poor state of preservation. The plants of the type material 
of M. variabile are a little larger and slightly more compact than 
the plants of M. tenerum. The angle of divergence of the branches, 
the flagelliform branches, the rhizoid production on the under- 
leaves, and the divergence and shape of the underleaves are identical 
(compare FIG. 5, nos. 1-4 and nos. 10-14). The cell walls are a 
little thinner but this difference is neither uniform nor constant. 
M. varibile is so similar to the parts of the plants of M. cuneis- 
tipulum with quadrate-orbicular underleaves that I have concluded 
that they belong to one species. Based on the available material, 



The portion of the type material available for study consists of one 
branched stem approximately 1.6 cm. long. 

FIG. 5. Bazzania cuneistipida (Gottsche & Lindenb.) Trevis. 1. Por- 
tion of a plant, ventral view, X 30. 2. Leaf, X 30. 3. Cells from the apical 
portion of a leaf, X 260. 4. A cell from the apical portion of a leaf, X 350. 
5. Portion of a plant, ventral view, X 30. 6. Apices of leaves, X 30. 7. A 
cell from the apical portion of a leaf, X 400. 8. Portion of a plant, ventral 
view, X 30. 9. A cell from the apical portion of a leaf, X 400. 10. Portion 
of a plant, dorsal view, X 30. 11. Portion of a plant, ventral view, X 30. 
12. Portion of a ventral leafy branch, X 30. 13. Cells from the apical portion 
of a leaf, X 260. 14. A cell from the apical region of a leaf, X 350. Nos. 
1-4 drawn from a portion of the type of M. variabile, from Puerto Rico; 5-7 
from the type of M. brevifolium, from Colombia; 8-9 from the type of M. 
corticola, from Brazil; 10-14 from the type of M. tenerum from St. Kitts. 



M. Fulford 24 Bazzania 

except for the trapezoidal outline of many of the underleaves of 
many of the plants of M. cuneistipulum, the two seem to be identical. 
Additional collections from all types of habitats will no doubt add 
to our knowledge of the variability of the species. None of the 
plants of M. tenerum or M. variabile had sexual branches. Ac- 
cording to HAMPE and GOTTSCHE (1852, 348), the female bracts 
of M. variabile are ovate, with the apices bifid to bidentate, and the 
lateral margins are dentate to long ciliate. This agrees with the 
female bracts of M. cuneistipulum. The female bracts of M. 
tenerum were not described. 

M. brevifolium G. from Colombia is another species with char- 
acteristics similar to those mentioned above. The type consists of 
a portion of a plant 1 cm. long, very poorly developed. The 
leaves at the growing point of the stem tend to be small and juvenile, 
and none of them are well developed. GOTTSCHE (1864, 141) has al- 
ready mentioned the similarity to B. longistipula (M. elegantulum) , 
and also to M. cuneistipulum, and since most of the leaves are 
bidentate one suspects that the species does not belong to the 
Tridentatae group. STEPHANIES figure (in the Tridentatae Sub- 
genus, in the I cones, Mastigobryum no. 153) shows the leaves to 
be two- or three-toothed, and the margins of the teeth serrate. He 
also describes the species in two places in the Species Hepaticarum 
(3 : 467 and 517) . These descriptions agree in most of their details, 
but in the earlier one he describes the leaves as tridentate (often 
bidentate), while in the later one he describes them as bidentate 
(rarely tridentate). The portion of the type from the STEPHANI 
Herbarium in Geneva consists of stems with most of the leaves 
bidentate (see FIG. 5, nos. 5 and 6). The margins of the teeth 
are entire. The species is another example of B. cuneistipula. 

M. corticola, collected in Brazil by WEINIO, should also be in- 
cluded here. The leaves are two-toothed, the cells are thin-walled 
but have conspicuous trigones, and the underleaves are subquadrate 
with convex sides (see FIG. 5, nos. 8-9). 

The species is readily distinguished from B. bidens and B. 
phyllobola because of their large, spreading leaves with large teeth, 
and from B. Herminieri because of its ovate, vittate leaves with 
large unequal teeth. 

DISTRIBUTION: Cuba: without locality, Wright, Hep. Cub. (H). 
Guadeloupe : without locality, THerminier (H). The specimen cited by 
Stephani (1903, 41) from Guadeloupe, collected by Duss, should be referred 
to B. Herminieri. Jamaica : without locality, Hb. Hooker, the type (V) ; 
without locality, Hart, cited by Boswell (1887, 50); New Haven Gap, Under- 
wood 911, 1024, 1054 (Y, NY); slopes of Sir John, E. G. Britton 1193 (Y); 
near Cinchona, Bower, cited by Pearson (1931, 96); without locality, Borgen- 
sen, Jansen, Harris, cited by Stephani (1908, 438). Martinique : Mt. 
Pelee, Duss 638 (H). Puerto Rico: without locality, Schwanecke, 
the type of M. variabile (B) ; Sierra de Naguabo 690-1035 m., Shafer, a, e, 
3712 (NY). St. Kitts : Mt. Misery, Breutel, the type of M. tenerum 
(V). Trinidad: without locality, Criiger, cited by Stephani (1908, 
438). Brazil : Lafayette, Weinio, the type of M. corticola (H). 
Colombia : Mt. Quindio, Triana and Planchon, the type of M. brevi- 
folium (G). 



Bazzania 25 Bidentatae 

REFERENCES: Boswell (1887, 50); Gottsche (1864, 141); Jack and 
Stephani (1892, 13); Lindenberg and Gottsche (1851, 70, 71. pi. 13); Pagan 
(1939, 39); Pearson (1931, 96); Stephani (1888a, 279; 1903, 23; 1908-09, 
437, 438, 467, 517; Icones, Mastigobryum nos. 27, 35, 36, 153, 155). 

5. Bazzania platystipula Fulford, Bryol. 44: 145-146. Fig. 1-11. 

1941. 

Plants growing in depressed mats or scattered among other 
bryophytes, light green becoming yellow-brown in the older por- 
tions ; the stems slender, often coarse thread-like, to 5 cm. long, with 
leaves to 2 mm. broad, prostrate ; stem cells in longitudinal section 
elongate, the medullary cells averaging 0.17 mm. long, the cortical 
cells shorter, deeply pigmented, both averaging 22ju in diameter, 
the end walls thin, the vertical walls uniformly thickened and con- 
taining frequent pits; lateral branches frequent, mostly 5 mm. 
apart, diverging at a wide angle ; flagelliform branches numerous, 
long ; rhizoids present on the leaves of the flagelliform branches : 
the line of leaf insertion curved in its upper part; the leaves ap- 
proximate to densely imbricated, ascendent, nearly plane, becoming 
strongly deflexed when dry, unsymmetrically ovate, 0.7 mm. - 1.2 
mm. long, 0.5 mm. broad at the base, narrowed to the obliquely 
truncate, bidentate apex ; the dorsal margin arched from a curved 
base, covering one-half the stem, the ventral margin straight, the 
base scarcely dilated, the apex two-toothed, the teeth acute to 
acuminate, two to four cells high, two to three cells broad at the 
base, the sinuses lunulate to acute, the margins entire; leaf cells 
large, thin-walled, the cell lumina angular-rounded, the trigones 
large, with convex sides, sometimes becoming confluent, the cuticle 
smooth to faintly verruculose ; cells of the apical portion and dorsal 
base 24/A - 32/t in diameter, of the median portion larger, and of the 
base 46/x - 50/A X 32/A, not forming a vitta : underleaves approximate 
to imbricated, subquadrate in outline, broader than the stem, the 
line of attachment straight, to 0.65 mm. long and broad, the lateral 
margins a little convex from a straight base, the apex undulate to 
deeply f our-lobed, the cells as in the apical portion of the leaf : 
leaves of the flagelliform branches scale-like, ovate, acute to 
shortly bidentate: female branches solitary, one to several on a 
stem, the bracts and bracteoles similar, ovate, the outermost series 
small, entire to shortly bidentate, the intermediate series larger, 
to one-third divided into usually three laciniae, the lateral margins 
ciliate to dentate; the innermost series (immature) to one-third 
divided into three laciniae, the lateral margins usually long ciliate 
and serrate to dentate: the perianth (immature) mouth short 
ciliate: male branches and sporophytes not seen. 

HABITAT: On tree bases and logs, in mats or scattered among 
other bryophytes. 

The distinguishing characteristics of the species are its small 
size, the greenish or yellowish brown color; the ascendent, shortly 
bidentate leaves, with large, thin-walled cells and large, rounded 
trigones with convex sides; and the approximate to imbricated, 
subquadrate underleaves with undulate to f our-lobed apices. (FlG. 

6. nos. 1-11). 




FIG. 6. Bazzania platystipula Fulford. 1, 2. Portion of a plant, 
ventral view, X 15. 3. Portion of a stem and leaf, dorsal view, X 30. 4. A 
tooth of a leaf, X 310. 5. A cell from the apical portion of a leaf, X 400. 
6, Underleaves, X 30. 7. Portion of the apical margin of an underleaf, X 310. 
8, Portion of a transverse section of a stem, X 310. 9. Cells of one of the 
laciniae of an innermost female bract, X 310. 10. Cells from the lower portion 
of the same bract. X 400. 11. Cells from the mouth of the perianth, X 310. 
Nos. 1-11 drawn irom the type material. 



Bazzania 27 Bidentatae 

The plants are slender, coarsely to finely filiform and form 
depressed mats on the bases of trees or logs. They vary in color 
from light green at the growing tips to yellow-brown below, some- 
times becoming very dark. The leaves are imbricated, ascendent, 
deflexed or even curled around the stem when dry, and are biden- 
tate with conspicuous, acute teeth. Some of the leaves on a few 
of the stems appeared to be caducous but the underleaves were 
persistent. 

The underleaves are large, conspicuously broader than the stem 
in robust plants (see FIG. 6, no. 1), where they are imbricated. 
On the less well developed stems they are not quite so broad, but 
nevertheless are large, and are usually approximate. The lateral 
margins are convex, entire, and the apices are undulate to two- to 
four-lobed (see FIG. 6, nos. 1, 2, 6). 

The cells of both the leaves and underleaves are very large (see 
FIG. 6, nos. 4, 5, 7), and have thin walls. The trigones are large 
and have convex sides so that they appear to be rounded. They 
form one of the conspicuous characteristics of the species. 

The female branches were poorly developed and in a very poor 
state of preservation. The intermediate and innermost series are 
to one-third divided into usually three laciniae (see FIG. 6, no. 9), 
and the lateral margins are variously incised, serrate, dentate and 
ciliate. The perianths were not developed except for the cells of 
the mouth. This appears to be short ciliate (see FIG. 6, no. 11). 

The species is similar in several respects to B. roraimensis, the 
description of which follows. It can be separated from the other 
Bidentatae of similar size and habit because of its large under- 
leaves, and the large, thin-walled cells with very large, rounded 
trigones. 

DISTRIBUTION : Ja m a i c a : John Crow Peak, 5500-5800 ft., Underwood 
692, the type (NY, Y) ; Blue Mountain Peak, Patterson 23 (F). Puerto 
Rico : Luquillo Mountains, E. G. Britton (NY). 

6. Bazzania roraimensis (Steph.) comb. nov. 
Mastigobryum roraimense Stephani, Trans. Linnean Soc. Bot. II. 6 : 97. 1901-05. 

Plants in mats, green to golden brown, becoming darker in the 
older parts ; stems slender to filiform, to 4 cm. in length, with leaves 
to 1.5 mm. broad, prostrate: lateral branches frequent, 5 mm. or 
more apart, diverging at a wide angle, sometimes becoming 
flagellif orm ; ventral branches flagellif orm, numerous, long ; rhizoids 
frequent, from the bases of the leaves of the flagellif orm branches: 
the line of leaf insertion curved in its upper part : leaves approxi- 
mate to imbricated, plane, becoming strongly deflexed when dry, 
ascendent, unsymmetrically narrow-ovate to oblong, 0.6 mm. - 0.8 
mm. long, mostly 0.42 mm. broad at the base, narrowed to 0.15 mm. 
at the shortly bidentate apex; the dorsal margin arched from a 
straight base, covering less than one-half the stem, the ventral 
margin straight, the base not dilated, the apex two-toothed, the 
teeth sharp, two to four cells high, two to four cells broad at the 
base, the sinuses lunulate, the margins entire; the leaf cells thin- 




FIG. 7. Bazzania roraimemis (Steph.) Fulford. 1. Portion of a plant, 
ventral view, X 30. 2. Portion of a stem with leaf, dorsal view, X 30. 3. A 
tooth of a leaf, X 310. 4. Cells from the apical portion of a leaf, X 400. Nos. 
1-4 drawn from a portion of the type material. 



Bazzania 29 Bidentatae 

walled, the cell lumina angular-rounded to stellate, the trigpnes 
conspicuous, large, often becoming confluent, the cuticle faintly 
verruculose ; cells of the apical region and dorsal base 20/x - 24/t in 
diameter, those of the median portion larger, those of the base to 
50/x X 20/x, a vitta not differentiated : underleaves distant, rounded 
in outline, not as broad as the stem, attached in a straight line, the 
margins entire, the apex sometimes undulate, the cells of the 
interior mostly 20/x in diameter, thin-walled and with conspicuous 
trigones, the marginal row thick-walled : leaves of the flagelliform 
branches scale-like, ovate, acute to shortly bidentate : male branches 
catkin-like, solitary, one to several on a stem, the bracteoles similar 
to the bracts, smaller, the bracts ovate, bidentate, the antheridia 
occurring singly: female branches and perianths not seen. (FlG. 
7, nos. 1-4). 

HABITAT : Not given. 

The portion of the type material in the STEPHANI Herbarium 
at Harvard does not agree in all of its characteristics with 
STEPHANI'S original description, or his figures in the Icones, 
Mastigobryum no. 34. His description and drawings were no doubt 
made from robust, well developed plants, while the plants in the 
Harvard collection are small and slender (see FIG. 7, nos. 1-4), 
with long internodes. The chief difference is in the underleaves. 
STEPHANI'S figures (Icones, Mastigobryum no. 34), and description 
show them as being broader than the stem, subquadrate in outline, 
and usually two- to four-lobed at the apex while the underleaves of 
the Harvard plants are small, scarcely broader than the stem, 
orbicular in outline, and entire or only faintly lobed along the apical 
margin. Unfortunately, it was not possible to obtain additional 
material so that a more complete examination could be made at 
this time. 

The plants (based on the Harvard material) are readily distin- 
guished from the other members of the Bidentatae by their brown 
color, the filiform stems with persistent leaves, the large cells with 
thin walls and very large trigones which soon become confluent, and 
the small, distant, orbicular underleaves which are usually entire. 

The species is similar to B. platystipula except for the size of 
the underleaves and the length of the internodes. Since, however, 
B. roraimensis, according to STEPHANI, does become more robust 
and the underleaves larger and more densely imbricated than the 
plants figured here, it may be that B. platystipula and B. roraimensis 
as described here, are different expressions of one species. 

Additional collections from Jamaica and British Guiana will no 
doubt furnish the necessary information but until further evidence 
should confirm this it seems best to consider the two as distinct. 
B. roraimensis can readily be separated from the small forms of 
B. cuneistipula because of its small, usually entire, orbicular under- 
leaves, and its large cells with thin walls and large trigones. 

DISTRIBUTION : British Guiana: summit of Mt. Roraima, McCon- 
nell and Quelch 523, the type (H). 

REFERENCES: Stephani (1908, 436; Icones, Mastigobryum no. 34). 



M. Fulford 30 Bazzania 

7. Bazzania gracilis (Hampe & Gottsche) Stephani, Hedwigia 27: 

279. 1888. 

Matttif/obryum gracile Hampe & Gottsche, Linnaea 25: 346. 1852. 
Mastiyobryum cuneifolium Gottsche, in Duss, Fl. Crypt. Antilles Fr. Hep. 23. 

1903. (nomen nudum) 

Mastiyobryum parvum Stephani, Spec. Hep. 3: 438. 1908. 
Bazzania bidenx var. vittata Spruce, Trans. & Proc. Bot. Soc. [Edinburgh] 

15: 371. 1885. 

Bazzania trichodcs Spruce ms. Hepat. Sprue. 
Monti yobryum trichoideum Stephani, Spec. Hep. 3: 532. 1909. 
Bazzania bidena Spruce, Trans. & Proc. Bot. Soc. [Edinburgh] 15: 371. 1885. 

Not M. bidem Gottsche & Lindenberg. 
Bazzania bidens var. Spruce ms. Hepat. Sprue. 

Plants scattered or growing in depressed mats, the slender 
forms yellow-brown to red-brown, larger plants yellow-green to 
olive-green; stems slender to filiform, prostrate to ascending near 
the growing tip, to 5 cm. long, with leaves 1.5 mm. - 2 mm. wide, 
leaves often becoming scale-like, on many of the stems caducous; 
stem cells in longitudinal section elongate, the medullary cells 
averaging 0.16 mm. long, the cortical cells shorter, both about 14/x 
in diameter, the end walls thin, the vertical walls irregularly thick- 
ened and containing pits; lateral branches frequent, 3 mm. - 10 
mm. apart, diverging at a very wide angle; flagelliform branches 
numerous, very long, often branched: the line of leaf insertion 
curved in the upper part; leaves distant to subimbricated, ascend- 
ent, nearly plane, becoming convex on drying, unsymmetrically 
ovate, becoming nearly symmetrical in the smaller forms; in the 
normal condition averaging 0.65 mm. - 0.85 mm. long X 0.4 mm. 
wide at the base, narrowing to the apex, often much smaller and 
scale-like, averaging 0.5 mm. or less long and 0.3 mm. wide at the 
base; the dorsal base straight to somewhat rounded, the dorsal 
margin convex in the typical forms, both the dorsal and ventral 
margins slightly convex in the reduced forms, the apex irregularly 
two-toothed becoming mostly acute in the smaller leaves ; the teeth 
two to three cells long, acute, the acroscopic tooth often the longer, 
the sinus broad, lunulate to acute; the leaf cells thin-walled, the 
trigones small, conspicuous, often confluent, the cell lumina 
angular-rounded, the cuticle smooth to faintly verruculose ; cells of 

FIG. 8. Bazzania gracilis (Hampe & Gottsche) Steph. 1. Portion of 
a stem of a small form, dorsal view, X 30. 2. Portion of a stem, ventral view, 
small form, X 30. 2a. Portion of a stem of large form, ventral view, X 30. 
3. Portion of stem with leaves much reduced, dorsal view, X 30. 4. Portion 
of stem of another large form, ventral view, X 30. 5. Leaves of the small 
form, X 30. 6. Tip of leaf of small form, X 260. 7. Cells from the dorsal 
margin of leaf of small form, X 260. 8. Basal cells of leaf (vitta) of small 
form, X 260. 9. Underleaf of the small form, X 30. 10. Cells from under- 
leaf of a small form, X 260. 11. Portion of a cross section of stem, X 260. 
12. Portion of a longitudinal section of a stem, X 260. 13. Male bracteole, 
X 30. 14. Male bracts, X 30. 15. Tip of male bract, X 260. 16. Female 
bract of outer series, X 30. 17. Female bracts of intermediate series (not 
mature) ? X 30. 18. Female bract of innermost series (not mature), X 30. 
19. Portion of perianth mouth (not mature), X 30. 20. Portion of perianth 
mouth, X 260. Nos. 1, 2, 2a, 5-12 drawn from a portion of the type material ; 
3. 13-15 from plants collected in Puerto Rico by EVANS, 57; 4, 16-20 from 
plants collected by UNDERWOOD, no. 659, in Jamaica. 



M. Fulford 32 Bazzania 

the apical portion and dorsal base 16/z - 20/i in diameter, the cells 
of the basal portion mostly 40/i X 15/i, forming a narrow but 
usually distinct vitta : underleaves small, distant, often as broad as 
the stem, attached in a straight line, on well developed stems often 
squarrose, quadrate to quadrate-orbicular, mostly 0.2 mm. - 0.32 
mm. long X 0.2 mm. - 0.3 mm. wide, much smaller in reduced 
forms, the lateral margins slightly convex, the apex entire, straight 
to rounded, or shortly one- to four-toothed or -lobed, the cells as 
in the apical portion of the leaf ; leaves of the flagellif orm branches 
scale-like, to 0.08 mm. long, ovate, entire to bidentate: male 
branches frequent, usually several on a stem ; the bracteoles small, 
oval, mostly 0.16 mm. X 0.08 mm., entire to bifid; the bracts 
larger, ovate, bifid, emarginate to crenulate, the cells of the apical 
portion averaging 27/x X 16/x, the cell walls thin, the trigones dis- 
tinct along the margins : female branches occasional, usually several 
on a stem: bracteoles and bracts similar, those of the outermost 
series small, quadrate, averaging 0.25 mm. X 0.08 mm., entire to 
bidentate; those of the intermediate series (not mature) ovate, the 
upper third or fourth divided into usually three, entire or dentate 
laciniae, the lateral margins often dentate; the innermost series 
(not mature), broadly ovate, the upper part divided into three 
laciniae, the margins dentate to ciliate, the cells all of one kind, 
elongate, mostly 45/i X 18^, thin-walled: perianth (not mature) 
mouth divided into numerous long laciniae similar to those of the 
innermost bracts. 

HABITAT: In depressed mats or scattered among mosses, tree 
bases, logs, and over rocks and soil. 

The species is easily distinguished from the other Bidentatae 
by its mostly slender or filiform appearance, and by the leaves 
which are small, often scale-like throughout much of the plant, 
two-toothed, with thin-walled cells, conspicuous trigones and a 
distinct vitta. They are usually caducous. The small form ap- 
pears as interwoven mats of brown hairs or threads while the 
larger forms have light green leaves. (FiGS. 8, nos. 1-20; 9, 
nos. 1-5). 

B. gracilis shows the usual wide series of variations from the 
large, green plants with persistent, well developed leaves and 
underleaves, to the filiform, red-brown stems usually without 
leaves except at the apices of the stems. A stem may produce 
normal, spreading leaves for a distance and then, through a gradual 
decrease in size, the small scale-leaves which are appressed, or it 
may produce only one kind. In the dry condition the branches 
with the small leaves appear similar to the flagelliform branches 
and only by a careful examination can they be distinguished. 
In the original description of the type material collected by 
SCHWANECKE, HAMPE and GOTTSCHE mention a variation p 
trichodes, which is depauperate, with the leaves squamiform except 
on shoots which tend to become erect, where they show a tendency 
to be large and spreading in the upper part of the stem. This 
description obviously has reference to plants in which most of the 
leaves are reduced, especially in the older portions of the plant. 




FIG. 9. Bazzania gracilis (Hampe & Gottsche) Steph. 1. Portion of 
a plant, ventral view, X 30. 2. A cell from the apical portion of a leaf, X 400. 
3. Portion of a plant, ventral view, X 30. 4. A cell from the apical portion 
of a leaf, X 400. 5* Portion of a plant, ventral view, X 30. Nos. 1-2 drawn 
from a portion of the type of M. parvum; 3-4 from a portion of the type of 
B. trichodes Spruce (NY) ; 5 from a portion of B. bidena Spruce (NY). 



M. Fulford 34 Bazzania 

The authors further state that the variation should not receive the 
dignity of a varietal rank. The cause of the alternating change in 
the kind of leaves produced is not known. It occurs in plants 
growing in a number of situations but is most conspicuous in the 
plants of higher elevations ; it may be due to the same influences 
which frequently cause the flagelliform branches to develop 
ordinary leaves. 

An interesting modification of the usual condition of the walls 
of the stem cells is often present. The thickenings of the vertical 
walls of the cells of the medulla, instead of being uniform, as is the 
usual condition, are very irregular, and in many of the cells small 
lumps of thickening protrude as warts into the cell-lumina as 
indicated in FIG. 8, no. 12. This same condition of irregular 
thickenings on the inner walls of the cells is present in the basal 
cells of the inner series of the female bracts and in the perianths 
of many of the West Indian species. 

Plants collected by UNDERWOOD in Jamaica were principally of 
the well developed type (FiG. 8, no. 4), described by HAMPE and 
GOTTSCHE (1852, 346) as the typical form. The stems are mostly 
3 cm. - 4 cm. long and with leaves mostly 1 mm. wide. The leaves 
are very often caducous and stems without leaves throughout much 
of their length are not uncommon. This caducous habit seems to 
be best developed in the normal type of leaf for in stems or parts 
of stems with reduced or scale-like leaves, rarely do they become 
detached. Furthermore, only rarely does one find this habit present 
in the underleaves, so that stems without leaves but with the 
underleaves persisting are not uncommon. 

The leaves are slightly ascendent, and this character becomes 
more strongly developed in the smaller forms (FiG. 8, nos. 1-4). 
They average 0.65 mm. - 0.85 mm. in length and 0.4 mm. in width 
and are unsymmetrically ovate. The dorsal margin is conspicu- 
ously arched from a straight to slightly rounded base ; the ventral 
margin is straight to a little concave. The apex is more or less 
transversely truncate and bidentate. The teeth show a high degree 
of variability in size. In most of the well developed leaves the 
teeth are mostly equal, two to three cells high, deltoid and acute, and 
separated by a lunulate sinus ; in the less well developed forms all 
variations are found from the sort just described, through the 
unequally two-toothed forms with a lunulate to deep and acute 
sinus, to finally, the sharply acute, scale-leaves of the filiform 
branches. 

The cells of the normal leaves are similar to those of the re- 
duced forms shown in FIG. 8, nos. 6-8, except that there are a 
greater number of elongated cells in the basal region. These ex- 
tend toward the middle of the leaf in one to three rows and form 
a fairly distinct vitta. 

The underleaves, FIG. 8, nos. 2 and 9, are small, quadrate, and 
appressed on the filiform stems but tend to become larger, FIG. 8, 
nos. 2a and 4, on the robust plants. Here they tend to be elongate- 



Bazzania 35 Bidentatae 

subquadrate, squarrose, and are approximately as broad as the 
stem. The lateral margins are mostly straight or only slightly 
bulging, usually mostly entire, and the apex is emarginate to 
two- to four-crenate or -lobed. 

The male branches were numerous on the plants collected by 
EVANS in the Luquillo Mountains, Puerto Rico. They were pro- 
duced on the normal stems as well as on the stems with reduced 
leaves. 

The type collection contains plants of two sorts ; larger plants as 
indicated in FIG. 8, no. 2a, with comparatively large leaves which 
are spreading, often caducous and have a more or less distinct 
vitta, and smaller plants with small, scale-like, persistent leaves 
which are appressed. The portion of the type material in the 
Farlow Herbarium (from the Hb. of JACK) is made up almost 
entirely of this smaller form, which HAMPE and GOTTSCHE distin- 
guish as /3 trichodes. 

The small red-brown variation seems to be more abundant in 
the West Indies than the large, leafy one, while in Central and 
South America the leafy variation is more often collected. 

The plants from Martinique distributed under the manuscript 
name M. cuneifolium Gottsche, and listed by Duss, are typical of 
the species. M. parvum collected by GLAZIOU in Rio de Janeiro is 
another example of the larger plants with caducous leaves (see 
FIG. 9, nos. 1-2). 

The plants described by SPRUCE from Mt. Chimborazo under 
B. bidens var. vittata which were distributed by him as B. trichodes 
and later described by STEPHANI as M. trichoideum also belong to 
B. gracilis. The plants are very similar to the larger plants from 
the West Indies except that they are not so strongly pigmented, 
the cell walls are not quite so thick, and trigones are not so large 
(FiG. 9, nos. 3, 4). Some of the leaves are caducous. B. bidens of 
SPRUCE from Rio Negro and Uaupes and his B. bidens var., "foliis 
brevioribus" from Manaos and San Carlos have the characteristics 
of his B. trichodes leaf shape, cell pattern, underleaves, etc., 
except that the plants are more robust, with the leaves more 
densely imbricated and larger (see FIG. 9, no. 5). The under- 
leaves are also larger but show the characteristic shape with the 
four-lobed apices. The leaves were not caducous on the plants 
examined. SPRUCE seems to have misinterpreted the species B. 
bidens, for he states that the figures of M. bidens in LlNDENBERG 
and GOTTSCHE, Species Hepaticarum, show the leaves a little 
longer than in his own specimens (which he considered typical), 
and that the form with long leaves was also collected in Guadeloupe 
by HUSNOT. I have examined the type of Herpetium stoloniferum 
var. bidens Mont, collected by LEPRIEUR in French Guiana and also 
the HUSNOT specimen and find no close similarity between them 
and SPRUCE'S plants. B. bidens can always be readily distinguished 
because of its long, more or less linear leaves. The cells are all 
very large and the trigones very large and often confluent. A vitta 
is not differentiated. 



M. Fulford 36 Bazzania 

DISTRIBUTION: Jamaica : Morce's Gap, 5000 ft, Underwood 530, 621, 
632 (Y, NY) ; base of John Crow Peak, 5000-6000 ft, Underwood 2361 (Y, 
NY) ; John Crow Peak, 5500-5800 ft, Underwood 657, 659, 823, 2360 ( Y, NY) ; 
Portland Gap, 1650 m., Maxon and Killip 1195 (H, Y, NY) ; New Haven Gap, 
Johnson 19 (Y) ; Cinchona, Mias Cummings 11 (NY) ; Green Hill Wood, 4000 
ft, Harris 12 (NY). Martinique: Mt. Pelee, Duss 638, as M. cunei- 
folium (NY). Puerto Rico: El Yunque, Luquillo Mountains, Evans 
57 (Y, NY); without locality, Schwanecke, the type (B, H); El Yunque, 
Pagan 486 (NY); without locality, Sintenis, cited by Stephani (1908, 436); 
Sierra de Naguabo, 465-720 m., J. A. Shafer 3758 (NY); Rio de Maricao, 
500-600 m., E. G. Britton 2682 (NY). Costa Rica: Without locality, 
Werckle (Y, NY); Zurqui, Standley 48091, reported as B. phyllobola by 
Herzog (W). Guatemala: Alta Verapaz near Coban, 1350 m., Tiirck- 
heim 5425 as M. phyllobolum (NY). Brasil : Rio de Janeiro, Glaziou 
(NY) ; Rio de Janeiro, Glaziou 4532, the type of M. parvum (H) ; Rio Negro 
and Uaup4s, Spruce, Hep. Sprue, as B. bidens (Y, NY) ; Manaos and San 
Carlos, Spruce, Hep. Sprue, as B. bidens var. (Y, NY). British 
Guiana : Mt. Roraima, McConnell and Quelch, 334/11 p.p., cited by Stephani 
(1901-05, 98). Ecuador : Mt Chimborazo, Spruce, Hep. Sprue., a por- 
tion of the type of B. trichodes Spruce (NY) ; with B. leptostipa, Spruce, Hep. 
Sprue. (NY). 

REFERENCES: Stephani (1888a, 279; 1901-05, 98; 1908, 436; /cones, 
Mastigobryum nos. 30, 32, 446) ; Pagan (1939, 39). 



SUBGENUS TRIDENTATAE 

This subgenus is characterized by having the leaves three- 
toothed or -lobed at the truncated apices. They are never or only 
occasionally two-toothed. Several species, namely, B. Schwaneck- 
iana, B. diversiciispis and B. canelensis, have most of the leaves 
entire or nearly so, but in each instance the species in question is 
very closely related to the species within one of the Sections 
listed below. 

The Sections Parvistipula* and Cordistipula-f of STEPHANI have 
been discarded because they were purely artificial groupings based 
on highly variable characteristics. 

The Section Connata of STEPHANI'S older classification (1886, 
245) has been revived to include all species in which the under- 
leaves are connate with one or both leaves. All of the American 
species of his Section Serrulata as well as one species from his 
Section Grandistipula belong here. 

Key to the Sections 

1. Leaves with a conspicuous vitta 5. Vittatae (p. 155) 

1. Leaves without a vitta. 

2. Underleaves connate with one or both leaves; leaf margins entire, 
inconspicuously serrulate to strongly dentate . . 2. Connatae (p. 103) 
2. Underleaves free from the leaves; leaf margins not serrulate. 

3. Underleaves divided to the middle into long lobes or teeth (no 
representatives reported for the Americas) 3. Fissistipulae (p. 124) 
3. Underleaves more or less entire, or less than one-third divided 
into lobes or teeth. 

4. Leaves with a conspicuous, usually appendiculate ventral 
auricle; Underleaves large, cordate, attached in- a recurved 
line; the line of attachment of the leaf curved in the upper 
part, the dorsal end bent downward forming a hook, 

4. Appendiculatae (p. 125) 

4. Auricles of the leaves, if conspicuous, never appendiculate; 
underleaves always attached in a straight line; the line of 
leaf insertion curved to hook-formed, 

1. Grandistipulae (p. 38) 



*The American species which STEPHANI included in the Section Parvi- 
stipula are included in the Grandistipulae section of this paper. Of the five 
species which he included in the Parvistipula, M. deciduum, M. elegantulum 
and M. Krugianum are small, depauperate, or juvenile forms of B. longi- 
stipula, p. 81. and M. tenue is a small plant of B. langa, p. 90. Since the small 
size of the plant, and the small, distant underleaves are the only criteria of 
the Section, it is often impossible to determine whether a specimen is an 
example of a species which is normally of small size, or is a juvenile or poorly 
developed form of a species of normally larger size, unless additional specific 
characteristics are taken into consideration. For this reason, M. tricuspidatum 
has also been transferred, and is treated under the Section Grandistipulae, 
p. 38. 

fAll of the American members of this Section have also been transferred 
to other Sections and are discussed elsewhere. 



Section 1. Grandistipulae 

Plants large or small, leaves without a vitta or conspicuous 
ventral auricles or appendages, the teeth with entire margins, the 
underleaves attached in a straight line, free from the leaves. 

Key to the Species 

1. Underleaves made up in part of hyaline cells which may form a con- 
spicuous border. 

2. Leaf cells uniformly thin-walled, trigones absent or minute, cell 
lumina rounded; underleaves hyaline except for a small internal area 

at the base 2. B. affinis (p. 39) 

2. Leaf cells thin-walled, trigones conspicuous, with bulging sides, cell 
lumina angular-rounded. 

3. Underleaves longer than broad, one-third to three-fourths hyaline, 

lateral margins scarcely bulging, never hyaline to the base; leaves 

elongate-ovate, the teeth large . . . 9. B. pallide-virens (p. 42) 

3. Underleaves subquadrate-rounded, the hyaline border continuous 

to the base. 

4. Hyaline cells thin-walled, chlorophyllose cells of the interior 

with conspicuous trigones . . . . 10. B. stolonifera (p. 44) 

4. Cell walls and trigones of the hyaline and chlorophyllose 

areas similar. 

5. Hyaline border always narrow, of one to four rows of 

cells 11. B. chilensis (p. 51) 

5. Hyaline border variable, absent, broad, or including most 

of the underleaf 12. B. taleana (p. 54) 

1. Underleaves chlorophyllose throughout. 

2. Underleaves variously toothed or deeply lobed or divided in the apical 
portion and on the lateral margins. 
3. Underleaves abundantly spinose-dentate and ciliate, 

13. B. denticulata (p. 56) 

3. Underleaves coarsely toothed or lobed but never ciliate. 
4. Underleaves deeply four-lobed in the apical portion, 

14. B. quadricrenata (p. 60) 

4. Underleaves variously toothed an-d lobed on the margins. 
5. Teeth of the leaves long, narrow, the margins crenu- 

late 15. B. aurescens (p. 63) 

5. Teeth of the leaves shorter, broad, the margins en- 
tire 16 B. Glaziovii (p. 65) 

2. Underleaves with entire margins, or only the apical margin retuse 
or with three or four undulations. 
3. Underleaves subquadrate. 

4. Plants large, green, underleaves often cordate. 

5. Lateral margins plane, entire, convex, the apex two or 
four-lobed or undulate . . . 17. B. Breuteliana (p. 68) 
5. Lateral margins recurved, convex, the apex retuse, 

18. B. acuminata (p. 75) 

4. Plants large, yellow-brown, underleaves cordate, attached in 
a recurved line (see Section Appendiculatae, B. Hookeri and 
B. robusta). 
4. Plants smaller, not as above. 

5. Teeth of the leaves very small or obscure, 

19. a diversicuspis (p. 77) 



Bazzania 39 Grandistipulae 

5. Teeth of the leaves well developed. 

6. Underleaves small, round, very distant on the stem; 
leaf apices broad, transversely truncate, trigones 

conspicuous* 20. B. tricuspidata (p. 79) 

6. Underleaves small to medium size, approximate to 
imbricated, broader than the stem. 
7. Leaves ascending, obliquely truncated, trigones 
conspicuous ... 21. B. longistipula (p. 81) 
7. Leaves spreading or falcate, teeth very coarse, 
cells and trigones very large, plants red- 
brown 23. B. longa (p. 90) 

7. Leaves spreading, cell walls thin, trigones in- 
conspicuous, underleaves often in part hyaline, 

12. B. taleana (p. 54) 

3. Underleaves longer than broad, rectangular in outline, the bases 
not cordate. 

4. Leaves ascendent, unequally three-toothed, cell walls thin, 
trigones conspicuous, plants green to yellow-brown, 

21. B. longistipula (p. 81) 
4. Leaves spreading to strongly falcate. 

5. Leaves spreading, long, teeth medium size; underleaves 
elongate, with parallel lateral margins; plants mostly 

golden yellow-brown 22. B. latidens (p. 88) 

5. Leaves falcate, sometimes spreading, unequally three- 
toothed, teeth large, coarse, cells and trigones very 
large; plants robust, yellow-brown to deep red-brown, 

23. B. longa (p. 90) 
3. Underleaves reniform, cordate at the bases. 

4. Underleaves inflated, appressed to the stem along the deeply 

pigmented margin, cells quadrate . 24. B. jamaicensis (p. 96) 

4. Underleaves plane, the margins not deeply pigmented, cells 

of the upper part elongate, with intermediate thickenings 

along the walls 25. B. Wrigiitii (p. 100) 

3. Underleaves ovate. 

4. Apical margin entire. 

5. Underleaves inflated, appressed to the stem along the 
deeply pigmented margin, cells quadrate, 

24. B. jamaicensin (p. 96) 

5. Underleaves plane, the margins not pigmented, cells of 
the upper part elongate, with intermediate thickenings 

along the walls 25. B. Wrightii (p. 100) 

4. Apical margin retuse, lateral margins recurved, 

18. B. acuminata (p. 75) 

8. Bazzania afflnis (Lindenb. & Gottsche) Trevis. Mem. 1st. Lomb, 

13: 415. 1877. 

Mastigobryum a fine Lindenberg & Gottsche, in G. L. & N., Syn. Hep. 720. 

1847. Not M. affine Mitten, in Hooker, Bot. Ant. Voy. 2': Fl. N. Zel. 

147. pi. C. fig. 4. 1854. 
Mastigobryum inciso-bilobatum Stephani, in Herzog, Biblioth. Bot. 87: 224. 

fig. 165, a-b. 1916. 

Plants scattered or growing in depressed mats, pale yellow- 
green to olive-green: stems slender, to 5 cm. or more in length, 
with leaves 2 mm. to 3 mm. broad, prostrate; in longitudinal sec- 
tion the cells elongate, the medullary cells averaging 0.18 mm. long, 

* Depauperate forms of other species might be confused with B. tricuspidata 
unless a careful examination of the plants is made. 



M. Fulford 40 Bazzania 

the cortical shorter, both averaging 0.18/x in diameter, the vertical 
walls uniformly thickened and containing frequent pits, the end 
walls thin ; lateral branches 0.5 cm. to 1.5 cm. apart, diverging at a 
wide angle; flagelliform branches frequent, long; rhizoids scarce, 
arising on the underleaves at the base : leaf insertion little curved 
in the upper part; leaves spreading, approximate to densely im- 
bricated, ascendent, becoming slightly falcate in robust plants, 
sometimes becoming deflexed when dry, unsymmetrically ovate, 
1.0 mm. - 1.6 mm. or more in length, mostly 0.56 mm. - 0.8 mm. 
broad at the base, narrowed a little to the transversely truncate, 
tridentate apex; the dorsal margin convex from a curved base 
which covers one-third to one-half the stem, the ventral margin 
convex, the base often more or less dilated ; the apex broad, more 
or less equally tridentate ; the teeth acute, deltoid, four to six cells 
long and broad, the sinuses lunulate to acute, the margins straight 
to undulate ; the leaf cells quadrate to rectangular in outline, with 
equally thickened walls, the cell lumina rounded, the trigones very 
small or absent; cells of the dorsal base and apical portion quad- 
rate, averaging 18/x X 18jn, those of the interior longer, averaging 
36/i - 45/x X 18/x, not forming a yitta; the cuticle verruculpse: 
underleaves distant to imbricated, in part hyaline, attached in a 
straight line, quadrate to oblong, mostly 0.48 mm. - 0.56 mm. long 
and wide, the apex straight to rounded, entire or with two to four 
short, broad teeth or lobes, the lateral margins nearly straight or 
bulging, entire or sometimes undulate, a little rounded at the 
bases; the cells with uniformly thickened walls and infrequent 
trigones, the hyaline cells occupying most of the leaf, the chloro- 
phyllose cells making up a small, internal basal area : leaves of the 
flagelliform branches small, ovate, the apex mostly rounded : sexual 
branches not seen. 

HABITAT : On decaying wood or on stones in woods. 

The distinguishing characteristics of the species are its dull, 
pale green prostrate stems, the tridentate leaves with quadrate cells 
and no or minute trigones ; and the entire or slightly two- to f our- 
lobed underleaves composed of hyaline cells throughout except for 
a small area of chlorophyllose cells in the basal region. (FiG. 10, 
nos. 1-12). 

The portion of the type of M. inciso-bilobatum from the Rijks- 
herbarium of Leiden was made up of plants identical with those 
of the type of M. affine from Mexico. Unfortunately, the illustra- 
tions which STEPHANI published with the original description of 
M. inciso-bilobatum and in the Icones, Mastigobryum no. 165 do 



FiG. 10. Bazzania affinis (Lindenb. & Gottsche) Trevis. 1. Portion 
of a plant, dorsal view, X 30. 2. Portion of a plant, ventral view, X 30. 
3. Leaf, X 30. 4. Apices of leaves, X 30. 5. A cell from the apical region of 
a leaf, X 350. 6. Cells from an apical tooth of a leaf, X 260. 7. Cells from 
the dorsal margin near the base, X 260. 8. Cells from the ventral margin 
including some from the basal portion, X 260. 9. Underleaves, X 30. 10. 
Portion of the base of an underleaf, including the margin, X 260. 11. Portion 
of the apical region of an underleaf, X 260. 12. Portion of a cross-section 
of a stem, X 260. Nos. 1-11 drawn from a portion of the type material; no. 
12 from material collected by A. C. SMITH in Fiji. 




pTO\ 

feramwo 1 

'M=~JL_.JL 



'. i:\.HJOOC 

V .iofecir ' 



ir 




M. Fulford 42 Bazzania 

not agree with these plants. His illustrations indicate that the 
underleaves are cordate at the base and bilobed at the rounded 
apex. The underleaves of the material from Leiden are quadrate, 
not cordate at the base, and have truncate, entire or undulate apical 
margins. 

DISTRIBUTION: Mexico: without locality or collector's name (NY). 
Colombia : Bogota, Andes, Weir (NY). Bolivia : Rio Tocorani, no. 
4073, Herzofc, the type of M. inciso-bilobatum (L). Peru : San Gavan, 
Lechler (NY). Fiji Islands: Koro; Vanua Levu: Thakaundrove, 
Smith (NY). 

REFERENCES: Lindenberg & Gottsche (1856, 93); Gottsche (1863, 164); 
Stephani (1886, 242'; 1908, 407; 1924, 468; loones, Maatigobryum nos. 151, 165). 

9. Bazzania pallide-virens (Steph.) comb. nov. 

Mastigobryum pallide-virens Stephani, Spec. Hep. 3: 473. 1908. 

Plants medium size, olive-green to dark green, becoming 
slightly pigmented with brown in the older portions : stems slender, 
to 4 cm. or more long, with leaves to 3.5 mm. broad; flagelliform 
branches numerous; rhizoids colorless, occasional on the bases of 
leaves of the flagelliform branches: the line of leaf insertion 
curved in its upper part ; leaves imbricated, mostly plane, unsym- 
metrical, ovate to elongate-oblong, straight to falcate, 1.8 mm. - 2.3 
mm. long, 5 mm. - 6 mm. broad at the base, narrowed a little to 
the transversely truncate, tridentate apex ; the dorsal margin con- 
vex from a curved base, covering one-half the stem, the ventral 
margins concave, the base little dilated, the apex sharply three- 
toothed, the teeth variable, three to eight cells long, three to six 
cells broad at the base, the sinuses deep, acute to lunulate, the 
margin entire; leaf cells thin- walled, the cell lumina angular- 
rounded, the trigones conspicuous, rarely becoming coalesced, the 
cuticle verruculose ; cells of the apical region 20/x - 32/A X 22/x, those 
of the median portion larger, those of the base 48/x - 60/A X 24ju, a 
vitta not differentiated : underleaves distant to subimbricated, ob- 
long, attached in a straight line, in part hyaline, the lateral margins 
nearly straight, entire, the apex truncate, entire, undulate or 
variously lobed, the hyaline border varying from several rows to 
the entire upper half of the leaf in width, the cells rectangular in 
outline, 24/i - 36/A X 20/i, those of the chlorophyllose area similar: 
the leaves of the flagelliform branches scale-like, ovate: male 
branches catkin-like, solitary, one to several on a stem, the bracts 
and bracteoles similar, concave, broadly ovate, shortly bifid, an- 
theridia occuring singly: female branches and perianths not seen. 

HABITAT : Not given. 

The distinguishing characteristics of the species are its green 
color; the mostly elongate, tridentate leaves with thin- walled cells 
and distinct trigones; and the oblong underleaves, one-third to 
three-fourths hyaline, with scarcely convex, entire, lateral margins 
and entire to variously lobed apices. (FlG. 11, nos. 1-6). 

The species differs from B. affinis in that its leaves are longer, 
the leaf cells have conspicuous trigones with bulging sides, and the 
basal portions of the underleaves are not bordered by hyaline cells. 




FIG. 11, Bazzania pallide-virens (Steph.) Fulford. 1. Portion of a 
plant, ventral view, X 15. 2. Portion of a leaf and stem, dorsal view, X 30. 
3. A leaf, X 15. 4. A tooth of a leaf, X 310. 5. A cell from the apical portion 
of a leaf, X 400. 6. Underleaves, X 30. Nos. 1-6 drawn from a portion of 
the type material. 



M. Fulford 44 Bazzania 

DISTRIBUTION: Brazil : without locality, Glaziou, 14418, the type (H). 
REFERENCES: Stephani (Icones, Mastigobryum no. 172). 

10. Bazzania stolonifera (Swartz) Trevis. Mem. 1st. Lomb. 13: 

415. 1877. 

Jungermannia stolonifera Swartz, Prodr. Fl. Ind. occ. 144. 1788. 
Pleuroschiama stolomiferum Dumortier, Receuil d'obs. Jungerm. 20. 1835. 
Herpetium stoloniferum Montagne in D'Orbigny, Voy. 1'Amer. Merid. T: 74. 

1839. 

Mastigobryum stoloniferum Lindenberg 1 in G. L. & N., Syn. Hep. 227. 1845. 
Jungermannia vincentiana Lehm. & Lindenb. in Lehman, Pug. PI. 4: 59. 1832. 
Herpetium vincentianum Mont. loc. cit. 

Mastigobryum vincentinum Lehm. & Lindenb. in G. L. & N., op. cit. p. 226. 
Bazzania vincentina Trevis. op. cit. p. 414. 

Mastigobryum Turkheimii Bvrd. in Stephani, Spec. Hep. 6: 481. 1924. 
Mastigobryum sylvaticum Stephani ms., non Gottsche, loc. cit. as synonym. 
Bazzania leptostipa Spruce, Trans. & Proc. Bot. Soc. [Edinburgh] 15: 374. 

1885. 

Mastigobryum leptostipum Stephani, Spec. Hep. 3: 524. 1909. 
Mastigobryum Lindigii Stephani, Hedwigia 25: 203. pi. 3. fig. 44-45. 1886. 
Bazzania Lindigii Spruce, Mem. Torrey Bot. Club 1: 129. 1890. 
Mastigobryum Quelchii Stephani, Spec. Hep. 3: 471. 1908. 

Plants large, dull olive-green to brownish green, light yellow- 
green at the growing tip: stems stout, to 10 cm. or more long, 
with leaves to 5 mm. wide, in deep tufts or depressed mats ; stem 
cells in longitudinal section averaging 0.16 mm. long, the marginal 
cells shorter, both containing pits, the end walls thin; lateral 
branches numerous, 1 cm. or more apart, diverging at a wide 
angle; flagelliform branches numerous, long, often branched; 
rhizoids colorless, present only on the leaves of the flagelliform 
branches ; the line of leaf insertion curved in the upper part ; leaves 
widely spreading, imbricated, nearly plane, becoming convex on 
drying, sometimes strongly deflexed, unsymmetrically ovate, to 
3 cm. long on robust plants, to 1.5 mm. wide at the base, narrowing 
to the truncate, tridentate apex, the dorsal margin strongly convex 
from a rounded base which covers two-thirds the stem, the ventral 
margin somewhat concave, sometimes dilated at the base, the apex 
transversely truncate, strongly to faintly tridentate, the teeth 
spreading, deltoid, acute to subobtuse, mostly unequal, large, three 
to seven cells long, three to seven cells broad at the base, the sinuses 
broad, lunulate, to acute, the margins straight to undulate; the 
cell cavities angular-rounded to stellate, the cell walls thin, the 
trigones small to large, with convex sides, often becoming confluent, 
the cuticle verruculose ; cells of the apical portion and dorsal base 
averaging 25/x long X 21/x wide, those of the median portion longer, 
those of the base mostly 45/A X 21/x, a vitta not differentiated: 
underleaves subimbricated to imbricated, attached in a straight 

FIG. 12. Bazzania stolonifera (Swartz) Trevis. 1. Portion of stem, 
dorsal view. X 12. 2. Portion of stem, ventral view, X 12. 3. Leaf, X 12. 
4. Apices 01 leaves, X 30, 5. A cell from the apical portion of a leaf, X 350. 
6. Apical tooth, X 260. 7. Cells from the dorsal margin of a leaf, X 260. 
8. Cells from the ventral margin of a leaf, X 260. 9. Cells from the basal 
portion of a leaf, X 260. 10. Underleaf, X 30. 11. Cells from the lateral 
margin of an underleaf, X 260. 12. Cells from the apical portion of an uncter- 
leaf, X 260. Nos. 1-12 drawn from the type, J. stolonifera, from Jamaica. 



M. Fulford 46 Bazzania 

line, broadly round-quadrate, mostly 0.7 mm. - 0.8 mm. long, 0.8 
mm. - 1 mm. wide, broader than the stem, with a hyaline border, 
the apex truncated, broadly undulate, slightly three- or four-lobed, 
the lateral margins convex, the bases rounded to semicordate ; the 
hyaline border continuous, four to eight or more cells wide, the 
cells more or less rectangular, thin-walled, 18//, - 30/x long X 
15/x - ISfjL wide, trigones rare; cells of the chlorophyllose internal 
portion averaging 25/i in diameter, longer near the line of attach- 
ment, the trigones with bulging sides as in the leaf, the cell lumina 
angular-rounded: leaves of the flagelliform branches ovate, 0.22 
mm. - 0.37 mm. long, squarrose-spreading, concave, the margins 
crenulate, the apex rounded to acute : male branches few to many 
on a stem, to 1 mm. long, ovoid, the bracteoles small, subquadrate, 
mostly 0.25 mm. long X 0.57 mm. wide, concave, mostly bilobed, 
the bracts larger, broadly ovate, mostly 0.6 mm. long X 0.5 mm. 
- 0.6 mm. wide, strongly concave, the apex bi- or tridentate with 
short, broad teeth, the lateral margins crenulate; the antheridia 
borne singly or in pairs: female branches several on a stem; the 
bracts and bracteoles similar, oblanceolate, the outer series 
averaging 0.67 mm. long X 0.45 mm. wide, the apex shortly bi- or 
tridentate, the innermost series larger, 1.5 mm. - 1.8 mm. long X 
0.85 mm. - 0.95 mm. wide at the base, the apex one-third to one- 
fourth divided into two to four long, narrow, crenulate teeth, the 
lateral margins crenulate, often undulate, the cells all of one kind, 
rectangular in outline, mostly 78/x X 20/A, cell walls, for the most 
part uniform, but with occasional protuberances on the inner walls 
of some of the cells: perianth to 6 mm. long, ovoid-cylindrical, 
three-keeled in the median portion, contracted at the short-ciliate 
to dentate mouth: capsule 1.5 mm. long, ovoid-cylindrical, of four 
or five layers of cells, the thickenings similar to those of B. 
trilobata; elaters to 390^, long, 18/x wide, bispiral, the ends rounded ; 
spores 15/x - 18/x, minutely punctate. 

HABITAT : In deep tufts or mats or mixed with other bryophytes, 
on soil, trees and logs in woods. 

The distinguishing characteristics of the species are its large 
size, the olive-green to brown-green color, the large, tridentate 
leaves with angular-rounded to stellate cell cavities and conspicuous 

FIG. 13. Bazzania stolonifera (Swartz) Trevis. 1. Portion of a stem, 
ventral view, X 10. 2. Stem and leaf, dorsal view, X 20. 3. Cells from the 
apical portion of a leaf, X 200. 4. A cell from the apical portion of a leaf, 
X 350. 5. Cells from the dorsal margin of a leaf, near the base, X 200. 
6. Cells from the ventral margin of a leaf near the base, including those from 
the middle of the leaf, X 200. 7. Cells from the lateral margin of an under- 
leaf, X 200. 8. Portion of a transverse section of a stem, X 230. 9. Apices 
of leaves, X 30. 10. Cells from the apical portion of a leaf, X 350. 11. Under- 
leaf, X 30. 12. Portion of a plant, ventral view, X 15. 13. A cell from the 
apical portion of a leaf, X 400. 14. A leaf, X 15. 15. Cells from the apical 
portion of a leaf, X 400. 16. Underleaves, X 15. 17. A cell from the apical 
portion of a leaf, X 400. 18. Underleaf, X 15. 19. A cell from the apical por- 
tion of a leaf, X 400. 20. Underleaf, X 15. 21. A cell from the apical portion 
of a leaf, X 400. 22. U.nderleaf, X 15. Nos. 1-8 drawn from plants collected 
by EVANS, no. 110, in Puerto Rico; 9-11 from the type of J. vmcentiama, from 
St. Vincent: 12-13 from the type of M< Lindigii, from Colombia; 14-16 from 
the type of B. leptostipa, from Ecuador; 17-18 from the type of M. Tiirkheimii, 
from Guatemala; 19-20 from the type of M. sylvaticum Steph., from Guate- 
mala; 21-22 from the type of M. Quelchii, from Guiana. 




SN^ 



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/ V{ ^ ^\ 

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15 




M. Fulford 48 Bazzania 

trigones; and the large, round-quadrate underleaves with con- 
tinuous hyaline borders of thin-walled cells surrounding the chloro- 
phyllose cells, the latter with conspicuous trigones. FlGS. 12, nos. 
1-12; 13, nos. 1-22; 14, nos. 1-10). 

The species is widespread throughout the West Indies and 
tropical South America, and because of its size and abundance is 
perhaps more frequently collected than most of the other species 
of this genus. As is the case of most of the other species, B. 
stolonifera also exhibits a wide variety of habitat modifications, 
particularly in the size of the trigones of the cells of the leaves, 
and in the width of the hyaline borders of the underleaves. 

The type specimens of J. stolonifera from Jamaica (see FIG. 12, 
nos. 1-12) are large, dark green plants deeply pigmented with 
brown, with large, round-quadrate underleaves, semicordate at the 
bases, and having a border of hyaline cells. The trigones of the 
leaf cells are very strongly developed, with bulging sides, and often 
become coalesced. The cell cavities are more or less stellate. 

In contrast to these are the plants of the type of J. vincentiana 
which agree in size, form, position of the leaves and underleaves, 
hyaline borders, etc., as is shown in FIG. 13, nos. 9-11, but differ 
in that they are less deeply pigmented and have a somewhat dif- 
ferent cell pattern. The cell walls are thin, and the trigones are 
small with bulging sides, and do not often become confluent, so 
that the cell cavity is angular-rounded rather than stellate. Be- 
cause the walls of the outer rows of cells are somewhat thicker 
than those of the internal cells, these cells appear to form a border 
around the leaf. While the marginal rows of cells of J. stolonifera 
also have thickened walls, the border is not conspicuous because of 
the large trigones and thick walls (coalesced trigones) throughout 
the leaf. This modification is not uncommon, and is illustrated 
further, FIG. 13, nos. 1-8, by plants from Puerto Rico. 

Plants of B. leptostipa collected by SPRUCE in Ecuador (FiG. 13, 
nos. 14-16), are somewhat smaller and more lax in habit and more 
delicate than the plants of J. stolonifera. The brown pigmentation 
is pronounced. On one stem the trigones of the cells of a leaf near 
the growing tip were conspicuous while those of the cells of leaves 
3 cm. farther down the stem were very much smaller. The under- 
leaves are distant to subimbricated. Many of them, especially 
those associated with leaves in which the trigones were minute, 
were mostly hyaline, with the chlorophyllose cells restricted to a 
small area near the base, but those on the upper part of the stem 
where the trigones of the leaves were larger had a much larger 
area of chlorophyllose cells, and were typical for the species. SPRUCE 
says of the leaves that they are deciduous. While several plants 
of the material examined were without leaves over a part of the 
stem, most of the plants still retained them, so that there may be a 
question as to whether these leaves were truly caducous or had 
become detached through some accidental means. All of the plants 
are very brittle when dry. This modification of the species at first 




FIG. 14. Bazzania stolonifera (Swartz) Trevis. 1. Male bracteole, 
X 40. 2. Male bracts, X 40. 3. Male bract with two antheridia, X 50. 4. Tip 
of male bract, X 100. 5. Female bract of outermost series, X 40. 6. Female 
bracts of an intermediate series, X 40. 7. Tip of female bract of inter- 
mediate series, X 200. 8-9. Female bracts of the innermost series, X 40. 
10. Tip of female bract of innermost series, X 200. Nos. 1-4 drawn from 
plants collected by UNDERWOOD, no. 825, on John Crow Peak, Jamaica; 5-10 
drawn from plants collected by EVANS, no. 110, at El Yunque, Puerto Rico. 



M. Fulford 50 Bazzania 

glance suggests B. affinis but a close examination will show that it 
does not belong to the latter, because of the trigone pattern of the 
cells of the leaves and the chlorophyllose cells of the underleaves 
of the more vigorous plants. The leaves of B. affinis have uni- 
formly thickened walls without trigones, or with very inconspicu- 
ous ones, and the underleaves are made up mostly of hyaline cells 
with the trigones not conspicuous in the green cells, 

A further degree of modification in this same direction is 
exhibited by plants of the type of M. Turkheimii and the ms. species 
M. sylvaticum of STEPHANI (FiG. 13, nos. 17-20), collected in 
Guatemala. These plants are light green in color, very large, the 
underleaves have very broad, hyaline borders, and the leaf cells are 
thin-walled, with very small trigones which have bulging sides. 
The walls of the marginal rows of cells are unthickened. Here 
again, the differences are a matter of slight degrees of modification 
and are probably due to a combination of habitat factors. 

M. Lindigii collected in Colombia is also a representative of the 
extremely mesophytic modification of the species. The plants are 
light green with some degree of brown pigmentation. The cells of 
the leaves have thin walls and very small trigones (FiG. 13, no. 13) . 
The underleaves are large and the hyaline borders well developed 
(FiG. 13, no. 12). 

The plants of M. Quelchii collected in British Guiana are robust 
and golden brown in color. The cells of the leaves have thin walls 
but the trigones are large and are very often confluent, and the 
underleaves are distinctive, with the usual typical hyaline borders 
as is shown in FIG. 13, nos. 21 and 22. 

The range of variation in trigone size seems to be due to a com- 
bination of the age of the plants and the habitat conditions, as has 
been shown by BUCK to be the condition in the genus Scapania. 
The degree of pigmentation also varies according to the habitat 
and age of the plant, as is the condition in other species of the genus. 

Male and female branches are frequently present. The female 
bracts and bracteoles and the perianth mouth are distinctive and 
will furnish further diagnostic characteristics (see FIG. 14, 
nos. 1-10). 

DISTRIBUTION : Cuba: without locality or collector's name, cited by 
Stephani (1909, 527); Santiago de Cuba, Funck & Schlim, 2097 (H). 
Dominica : Morne Micotrin, 97, 99, Morne Couronne, 218, Elliott, cited 
by Spruce (1895, 356). Guadeloupe: without locality, Madianna 
(NY); without locality, 1'Herminier 55, 119 (NY); without locality, Huanot, 
PL Antilles, 212 (NY); Mornes des Deus Marnelles, Duss 282 p.p. (NY); 
nos. 117, 282 p.p. from Morne Hirondelle are B. Breuteliana; without locality, 
1'Herminier, Gottsche & Rabenhorst, Hepat. eur. 561 (Y, NY) ; without 
locality, Grateloup, Hb. Montagne, cited by Bescherelle (1893, 186); St. Rose, 
1'Herminier (H); Soufriere, Duss 66, 205, 320 as M. portoricense (NY). 
Jamaica: without locality, Swartz, the type (B, NY, H) ; without locality, 
Menzies 101 (NY); without locality, Wilson (NY); without locality, Hart, 
cited by Boswell (1887, 50) ; John Crow Peak, near Cinchona, E. G. Britton 
213 (NY); base of John Crow Peak, 5000-5500 ft, Underwood 2327, 2328, 
2370, 2421, 2426 (Y, NY) ; John Crow Peak, 5500-5800 ft., Underwood 684, 
825, 832, 834, 2427, 0428 (Y, NY) ; Morce's Gap, 1500 m., Maxon & Killip 



Bazzania 51 Grandistipulae 

647 (Y, NY); Blue Mountain, Bennet (NY); Britton 1103, 1190, 1227 (Y, 
NY) ; Greenwich Woodlands, 4500 ft., Harris 11,170 (NY) ; New Haven Gap, 
Evans 537 (Y) ; near Hardware Gap, 4000 ft., Underwood 2240 (Y, NY); 
Doll Woods, Evans 495 (Y) ; road from Cinchona to Morce's Gap, 5000 ft., 
Underwood 273 (NY) ; without locality, Wilson 768 (NY) ; Morce's Gap, 
New Haven Gap, Patterson (F). Martinique : without locality, Hus- 
not, PI. Antilles, 212 (NY) ; Calebasse, Mt. Pelee, Duss 13, cited by Stephani 
(1903, 22), 117 (NY). Puerto Rico: El Yunque, Evans 110 (NY); 
Sierra Luquillo, Br. Hioram 403 p.p. (NY) ; without locality, Sintenis, 16, 
cited by Stephani (1888a, 279); without locality, Sintenis 92, as B. Wrightii 
(NY) ; wthout locality, Schwanecke, cited by Hampe & Gottsche (1852, 350) ; 
Mt. Britton, Jones 10978 (H). St. Kitts : without locality, Breutel 
(NY); Mt. Misery, Breutel (NY); Britton & Cowell 508 (NY); without 
locality or collector, cited by Stephani (1909, 527). St. Vincent: 
without locality, Hb. Hooker 170 (NY) ; St. Andrews, Guilding, Hb. Hooker, 
the type of J. vincentiana (NY) ; without locality, Menzies (NY) ; without 
locality, Hart, cited by Boswell (1887, 50); St. Andrews, Elliott 63, cited by 
Spruce (1895, 356); without locality, G. W. Smith (H). Mexico: 
ft intermedia, Vera Cruz, Liebmann 52, cited by Gottsche (1863, 231). 
British Honduras : Camayaguam, 1800 m., Yuncker, Dawson & Youse, 
6617, 6618 (F). Costa Rica : Cartago, Maxon 502 (NY); Yerba 
Buena, Standley 49949, 49882, 49842; forma defolians 49869; forma minor, 
ramulosa 49891; Laguna de la Chonta, Standley 42240; Alto de la Estrella, 
Standley 39101; Cerro de las Lajas, Standley 51612 f. rufescem (W). 
Guatemala: Coban, Turkheim 5816, the type of M. Turkheimii (G) ; 
no. 5582 as M. viridissimum, no. 5418 as M. Quelchii (H). Nicaragua : 
Segovia (Ocotal), Oersted, cited by Hampe (1851, 302). Bolivia : with- 
out locality, D'Orbigny, cited by Montagne (1839, 74); also H, vincentianum 
from the same reference; Corana, Herzog 3390a, cited by Stephani (1916, 
225) as M. Lindigii. - B r a z i 1 : Rio de Janeiro, Glaziou 4572 (NY); the 
same, cited by Nees von Esenbeck in Martius (1833, 1: 337) ; Espirito, Santo, 
Ynes Mexia 4076 (H, NY). Colombia : Merida, Moritz, cited by Hampe 
(1847, 328); without locality, Lindig, the type of ,Af. Lindigii (H); without 
locality, Wallace (NY); var. granatensis cited by Gottsche (1864, 140). 
Ecuador : Mt. Tunguragua, Spruce (1885, 377); Mt. Chimboraza, Spruce, 
Hepat. Sprue., the type of B. leptostipa (Y). Guiana : var. irregulare 
Nees et Mont, ms., Leprieur 292, cited by Montagne (1840, 333). B r i t i s h 
Guiana: Mt. Roraima, McConnell & Quelch, cited by Stephani (1901-05, 
98); Mt. Roraima, Quelch, the type of M. Quelchii (H). Listed also from 
the Island of Bourbon, Bory; Australia and Sandwich Islands. 

REFERENCES: Swartz (1806, 3: 1862) ; Weber & Mohr (1807, 409) ; Lunan 
(1814, 518); Schwaegrichen (1814, 19); Weber (1815, 43); Sprengel (1827, 
4: 222); Nees von Esenbeck (1830, 61; 1833b, 376); Wallroth (1831, 75); 
Montagne (1840, 333; 1844-46, 242; 1855, 315); G. L. & N. (1844-47, 720); 
Hampe (1847, 328; 1851, 302; 1873, 227); Lindenberg & Gottsche (1851, 71, 
78); Hampe & Gottsche (1852, 350); Gottsche (1863, 231; 1864, 140); Husnot 
(1875, 3) ; Spruce (1885, 377; 1895, 356) ; Boswell (1887, 50) ; Stephani (1888a, 
279; 1901-05, 98; 1903, 22; 1908, 471; 1909, 526; 1924, 481; /cones, Mastigo- 
bryum nos. 168, 174, 176, 178, 391, 406) ; Bescherelle (1893, 186) ; Herzog (1938, 
19); Pagan (1939, 39). 

11. Bazzania chilensis (Steph.) comb. nov. 

Mastigobryum chilense Stephani, Hedwigia 24: 247. pi. 2. fig. 1. 1885. 

Plants medium size, dark green, pigmented with brown : stems 
to 3 cm. (?) long, with leaves to 3 mm. broad; lateral branches 
frequent, diverging at a wide angle ; flagellif orm branches frequent ; 
rhizoids colorless, present on the leaves of the flagellif orm branches : 



M. Fulford 52 Bazzania 

the line of leaf insertion curved in its upper part ; leaves imbricated, 
plane, becoming a little deflexed when dry, unsymmetrically ovate, 
straight or nearly so, 1.4 mm. - 1.8 mm. long, 0.5 mm. broad at 
the base, narrowed to the transversely truncate, tridentate apex; 
the dorsal margin strongly arched from a rounded base, covering 
two-thirds of the stem, the ventral margin straight, the base 
scarcely dilated, the apex three-toothed, the teeth large, eight to 
twelve cells long, five to ten cells broad at the base, the sinuses 
deep, acute to lunulate, the margins entire; the leaf cells thin- 
walled, the trigones conspicuous, often confluent, the cell lumina 
angular-rounded, the cuticle strongly verruculose; cells of the 
apical portion mostly 20/x X 20/x, of the median portion larger, 
and of the base 40/z X 22/1, a vitta not differentiated : underleaves 
distant to imbricated, attached in a straight line, broader than the 
stem, round-quadrate in outline, mostly 0.43 mm. long and 0.5 mm. 
broad, with a hyaline border, the lateral margins convex from a 
rounded base, the apex truncate, mostly straight, crenulate, the 
hyaline border of one or two rows of cells on the lateral margins, 
of two to four rows at the apex, the cells averaging 16/4. in diameter, 
the walls thin, the trigones small but distinct, those of the chloro- 
phyllose area similar: sexual branches not seen. 

HABITAT: Not given. 

The distinguishing characteristics of the species are its medium 
size, the dark green color ; the strongly tridentate leaves with small, 
thin-walled cells with conspicuous trigones ; and the round-quadrate 
underleaves with a border of one to three rows of hyaline cells not 
distinct from the chlorophyllose cells in size, wall thickness, or 
trigone structure. (FiG. 15, nos. 1-6). 

The plants remind one of small forms of B. stolonifera but they 
differ from that species in several characteristics. The cells of the 
leaf are smaller, mostly 20/x X 20/x, while in B. stolonifera they 
are mostly 24/* - 30/ A X 24/x. The cuticle is more strongly verru- 
culose. The greatest difference between the two species is to be 
found in the underleaves. In B. chilensis the underleaves are 
smaller and the hyaline margin is not nearly so distinct. This 
hyaline margin varies from one to four cells in width in the apical 
portion, and is only one or two cells wide along the lateral margins 
as is seen in FIG. 15, nos. 1, 5 and 6. Only rarely is it entirely 
absent from the basal portion of the margin. These hyaline cells 
do not differ from those of the chlorophyllose interior except in the 
absence of chlorophyll (FiG. 15, no. 16), The cell walls and 
trigones are similar in the two areas. This characteristic alone is 
sufficiently striking to immediately separate B. chilensis from B. 
stolonifera, for in the latter the cells of the hyaline part are very 
different from those of the chlorophyllose area. 

STEPHANI listed several specimens, "Peru, Callao (N6LLNER), 
Tatanara (LECHLER), Chile (LECHLER), Nova Granada (SCHLIM 
861)" in his original description of the species, and did not desig- 
nate any one of them as the type. The plants collected by SCHLIM 
no. 861 in Colombia were the only ones available for study (see 
FIG. 15, nos. 1-6). They do not agree with the original description 




FIG. 15. Bazzania chilensis (Steph.) Fulford. 1. Portion of a plant, 
ventral view, X 15. 2. Portion of a leaf and stem, dorsal view, X 30. 
3. Apical tooth of a leaf, X 310. 4. A cell from the apical portion of a leaf, 
X 400. 5. Underleaf, X 30. 6. Cells from the apical portion of an under- 
leaf showing the hyaline margin, X 310. Nos. 1-6 drawn from a portion 
of the type (?) material, collected by SCHLIM in Colombia. 



M. Fulford 54 Bazzania 

and figures in Hedwigia or with the figures in the unpublished 
Icones, Mastigobryum no. 381, which were made from plants col- 
lected by LECHLER in Chile. It may be that a study of the other 
collections cited will bring to light plants of the sort illustrated by 
STEPHANI. If that is the case, the plants described above represent 
an up to now undescribed species. STEPHANI'S illustrations 
suggest robust plants of B. Breuteliana. 

DISTRIBUTION: Bolivia : Yungas, Rusby, cited by Spruce (1890, 129). 
Chile: without locality, Lechler, cited by Stephani (1885, 247). 
Colombia: without locality, Schlim 861 (H). Peru: Callao, 
Nollner, cited by Stephani (1885, 247); Tatanara, Lechler, cited by Stephani 
(1885, 247). 

REFERENCES: Stephani (1909, 522; Icones, Mastigobryum no. 381). 

12. Bazzania taleana (Gottsche) comb. nov. 

Mastigobryum taleanum Gottsche, Mex. Leberm. 131. 1863. 
Mastigobryum longiscuspe Stephani, Spec. Hep. 3: 472. 1908. 
Maxtigobryum variedentatum Stephani, in Herzog, Biblioth. Bot. 87: 225. 
Fi%. 166. 1916. 

Plants delicate, medium size, olive-green, becoming pigmented 
with brown in the older portions : stems to 3 cm. or more in length, 
with leaves to 2.5 mm. broad : lateral branches frequent, mostly 5 
cm. apart, diverging at a wide angle: flagelliform branches fre- 
quent; rhizoids not seen: the line of leaf insertion curved in its 
upper part; leaves imbricated, plane to deflexed, unsymmetrically 
ovate, ascendent, 1 mm. - 1.5 mm. long, 0.5 mm. broad at the base, 
narrowed a little to the transversely truncate, tridentate apex ; the 
dorsal margin convex from a curved base, covering one-half the 
stem, the ventral margin straight, the base scarcely dilated, the 
apex more or less equally three-toothed, the teeth narrow, sharp, 
five to eight cells long, four to six cells broad at the base, usually 
ending in a row of two cells, the sinuses broad, lunulate, the 
margins mostly entire ; the leaf cells quadrate in outline, the walls 
thin, the trigones minute, the cell lumina rounded, the cuticle faintly 
verruculose ; the cells of the apical region mostly 16/x - 20/x X 18ju, 
of the median portion larger, and of the base to 40/x X 24/x, a vitta 
not differentiated: the underleaves distant to imbricated, round- 
quadrate, attached in a straight line, broader than the stem, .55 
mm. - .65 mm. long X .55 mm. - .6 mm. broad, the lateral margins 
entire, convex from a rounded base, the apex mostly entire, repand 
to undulate, the cells as in the apical portion of the leaf, some- 
times hyaline in part: leaves of the flagelliform branches scale- 
like, ovate, acute to bifid : sexual branches not seen. 

HABITAT : Not given. 

The distinguishing characteristics of the species are its medium 
size and green color; the slightly ascendent leaves, transversely 
truncate, and set with three, nearly equal, short teeth often ending 
in two-celled points; the quadrate leaf cells with thin walls, 
minute trigones and rounded cell lumina; and the round-quadrate, 
repand underleaves with entire margins. The underleaves may be 
in part or entirely devoid of chlorophyll. (FlG. 16, nos. 1-10). 







lOv 



FIG. 16. Bazzania taleana (Gottsche) Fulford. 1. Portion of a plant, 
ventral view, X 30. 2. A cell from the apical portion of a leaf, X 400. 
3. Underleaf, X 30. 4. Portion of a plant, ventral view, X 15. 5. Portion of 
a leaf and stem, dorsal view, X 30. 6. A leaf, X 30. 7. A tooth of a leaf, 
X 310. 8. A cell from the apical portion of a leaf, X 400. 9. Underleaves, 
X 30. 10. A portion of the apical margin of an underleaf, X 310, Nos. 1-3 
drawn from a portion of the type of M. taleanum (H); 4-10 from a portion 
of the type of M. longicuspe (H). 



ML Fulford 56 Bazzania 

The plants of M. longicuspe from Brazil seem to be better de- 
veloped than those of M. taleanum from Mexico, although all of 
the material is fragmentary (see FIG. 16, nos. 1 and 4). The 
two-celled points of the teeth of the leaves (FiG. 16, no. 7), seem to 
be a constant characteristic on the more robust plants. 

The underleaves are round-quadrate, with the apex more or 
less retuse, and the margins entire, as is shown in FIG. 16, nos. 1, 
3, 4, and 9. It is not unusual to find an underleaf one-half to two- 
thirds hyaline adjacent to an underleaf which is chlorophyllose 
throughout its entire area. Since not all of the underleaves on a 
stem are completely or in part hyaline, it would seem that for this 
species the hyaline characteristic may be due to external conditions. 
There is no differentiation in structure between the hyaline and 
chlorophyllose cells. The margins of the underleaves of poorly 
developed plants are in part crenulate or even occasionally dentate. 

The plants of M. variedentatum collected by HERZOG in Bolivia 
are like those mentioned above except that none of the underleaves 
are hyaline. However, since this is a variable characteristic in 
both M. taleanum and M. longicuspe, there seems to be no justifica- 
tion for considering M. variedentatum a distinct species. B. taleana 
is readily distinguished from the other members of the Grandi- 
stipulae which have the underleaves in part hyaline, since the 
underleaves are retuse with entire margins, the hyaline areas of 
the underleaves of a stem vary in size and position, and the hyaline 
cells are not structurally distinct. 

DISTRIBUTION: Mexico: Oaxaca, Mt. Talea, Liebmann, the type (H). 
The portion of the type from the Herb. Boissier was a poorly preserved plant 
of some other species. Brazil : Apiahy, Puiggari, the type of M. 
longicuspe (H). Bolivia : Comarapa, Herzog, the type of M. varied- 
entatum (L). 

REFERENCES: Stephani (1909, 519: 1924, 468; I cones, Mastigobryum nos. 
169, 180, 401). 

13. Bazzania denticulata (Lindenb. & Gottsche) Trevis. Mem. 1st. 

Lomb. 13: 414. 1877. 

Mastigobryum denticulatum Lindenberg & Gottsche, in G. L. & N. Syn. Hep. 

718. 1847. 

Mastigobryum planiusculum Lindenberg & Gottsche, loc. cit. 
Bazzania planiuscula Trevis. loc. cit. 
Bazzania Rusbyi Spruce, Mem. Torrey Bot. Club 1 : 129. 1890. 

Plants medium size, light green becoming slightly pigmented 
with brown in the older portions : stems stout, to 5 cm. or more in 

FIG. 17. Bazzania denticulata (Lindenb. & Gottsche) Trevis. 1. Por- 
tion of a plant, ventral view, X 10. 2. Portion of a leaf and stem, dorsal 
view, X 25. 3. Leaf, X 25. 4. Cells from the dorsal base, X 300. 5. A cell 
from the apical portion of a leaf, X 400. 6. Underleaves, X 25. 7. Portion 
of the apical area of an underleaf, X 300.x 8. Female bracts of an outer 
series, X 40. 9. Female bract of an intermediate series, X 40. 10. Portion 
of a lacinia of this bract, X 100. 11. Female bract of the innermost series, 
X 40. 12. Portion of a lacinia of this bract, X 400. 13. Portion of a trans- 
verse section of a stem, X 400. Drawn from plants collected by MAXON and 
KILLIP in Jamaica. 




12 



M. Fulford 58 Bazzania 

length, with leaves to 3.5 mm. broad, prostrate ; stem cells in longi- 
tudinal section averaging 0.17 mm. long, the cortical shorter, both 
averaging 20/i in diameter, the vertical walls uniformly thickened, 
containing frequent pits, the end walls thin ; lateral branches fre- 
quent, 5 mm. or more apart, diverging at a wide angle ; flagellif orm 
branches numerous, short; rhizoids present on the leaves of 
flagelliform branches : the line of leaf insertion curved in its upper 
part, the leaves imbricated, plane, becoming a little convex on dry- 
ing, unsymmetrically ovate, straight, 1.5 mm. - 2 mm. long, to 
1 mm. broad at the base, narrowing to the more or less obliquely 
truncate, strongly tridentate apex ; the dorsal margin strongly con- 
vex from a rounded base, covering one-half or more of the stem, 
the ventral margin concave, the base sometimes dilated, the apex 
equally three-toothed, the teeth spreading, large, six to eight cells 
long, five to seven cells broad at the base, the margins entire, the 
sinuses acute to rounded, deep; leaf cells uniformly thin-walled, 
trigones scarcely evident, the cell lumina rounded, the cuticle 
faintly verruculose ; cells of the apical region averaging 20/x, of the 
median portion larger, and of the base 30/x - 40^ X 24/^, a vitta 
not differentiated: underleaves approximate to imbricated, squar- 
rose, subquadrate, broader than the stem, attached in a straight 
line, not connate with the leaves, 0.5 mm. - 0.7 mm. long X 0.6 
mm. - 0.9 mm. broad, the apex and lateral margins variously 
toothed with numerous long, narrow, often ciliate teeth or spines, 
becoming crenulate near the curved bases, the cells as in the leaves : 
leaves of the flagelliform branches scale-like, narrowly ovate, to 
0.3 mm. long, the apex bi-tridentate : female branches occasional, 
solitary, one to several on a stem, the bracts and bracteoles similar, 
the outermost series ovate-lanceolate, 0.65 mm. long X 0.45 mm. 
broad, the margins crenulate, occasionally toothed, the apex with 
two to four sharp teeth ; the intermediate series larger, to 2 mm. 
or more long, ovate-lanceolate, the margins ciliate to dentate, the 
apex divided into two to four sharp teeth or dentate laciniae; the 
innermost series similar, larger, more deeply divided into usually 
four dentate-ciliate laciniae, the lateral margins dentate to short 
ciliate, the cells rectangular in outline, 52^ - 90/x X 26/A, thin-walled : 
male branches and perianths not seen. 

HABITAT : Over mosses, tree bases and logs in wooded areas. 

The distinguishing characteristics of the species are its pale 
green color, the prostrate habit, the spreading, strongly dentate 
leaves with uniformly thin-walled cells and minute trigones; and 
the subquadrate underleaves with irregularly and variously spinose 
to ciliate, toothed margins. The underleaves are not connate with 
the leaves. (FIGS. 17, nos. 1-13; 18, nos. 1-6). 

The plants of the type material of M. planiusculum are not 
quite so large and do not have so many ciliate teeth on the margins 
of the underleaves as are to be found in M. denticulatnm, but these 
differences are not constant. The plants of B. Rusbyi are lighter 
green, larger, and often have more teeth on the underleaves than 
do those mentioned above. STEPHANI considered both M. planius- 
culum and B. Rusbyi identical with M. denticulatum and reduced 
them to synonomy. SPRUCE considered B. Rusbyi very close to the 




FIG. 18. Bazzania denticulata (Lindenb. & Gottsche) Trevis. 1. Por- 
tion of a plant, ventral view, X 15. 2. Portion of a leaf and stem, dorsal 
view, X 30. 3. A cell from the apical portion of a leaf, X 400. 4. Portion of 
a plant, ventral view, X 15. 5. A cell from the apical portion of a leaf, 
X 400. 6. Underleaf, X 30. Nos. 1-3 drawn from a portion of the type of 
M. planiusculum from Mexico; 4-6 from the type of B. Rusbyi from Bolivia. 



M. Pulford 60 Bazzania 

Mexican M. denticulatum, and states (1890, 116), that the latter 
"is possibly its nearest congener, but is smaller, more opaque, and 
the underleaves are much less toothed, especially at the sides." 
There seems to be no adequate reason for keeping them distinct. 

The species is easily distinguished from the other members of 
the Tridentatae because of its strongly three-toothed leaves, the 
uniform cell walls with inconspicuous trigones, and the ciliate, 
dentate underleaves. In appearance it resembles certain members 
of the West Indian Connatae Section, but the attachment of the 
underleaf, free from the leaves will at once distinguish it. 

DISTRIBUTION: Jamaica: John Crow Peak, Maxon & Killip 989 (NY); 
Morce's Gap, Evans 29 (Y); Cinchona, Earle (Y). Mexico: Oaxaca, 
Liebmann 298, the type (H) ; Oaxaca, Liebmann 295b, the type of M. planius- 
culum (G). B o 1 i v i a : Yungas, Rusby 3029, the type of B. Rusbyi (NY) ; 
also no. 5029 from the same locality. 

REFERENCES: Lindenberg & Gottsche (1851, 62, 63); Gottsche (1863, 136, 
137) ; Stephani (1886, 243; 1908, 506; Icones, Mastigobryum 345 a, b). 

14. Bazzania quadricrenata (Gottsche) Pagan. Bryologist 42: 39. 

1939. 

Mastigobryum quadricrenatum Gottsche in Stephani, Hedwigia 25: 206. pi. 1, 

fig. 1-4. 1886. 
Mastigobryum quadricrenatum forma paupercula G. A. Lindberg, Hedwigia 

25: 206. 1886. (nomen nudum). 
Mastigobryum Martianum Gottsche in Stephani, Hedwigia 25: 205. pi. 1, fig. 

9-12. 1886. 

Plants medium size to large, olive-green to brownish green, be- 
coming deeply pigmented with brown in the older portions : stems 
to 5 cm. or more long, with leaves to 3.5 mm. broad, prostrate to 
suberect: stem cells in longitudinal section 0.17 mm. long, the 
cortical shorter, averaging 20/* in diameter, the vertical walls uni- 
formly thickened and containing frequent pits, the end walls thin : 
lateral branches frequent, mostly 5 mm. apart, diverging at a wide 
angle; flagelliform branches frequent; rhizoids colorless, on the 
bases of leaves of the flagelliform branches : leaf insertion curved 
in its upper part ; the leaves imbricated, plane to deflexed, unsym- 
metrically ovate to oblong-ovate, nearly straight, 1.5 mm. - 2 mm. 



FIG. 19. Bazzania quadricrenata (Gottsche) Pagan. 1. Portion of a 
stem, ventral view, X 12. 2. Portion of a stem with leaf, dorsal view, X 30. 
3. A cell from the apical portion of a leaf, X 400. 4. An apical tooth of a 
leaf, X 310. 5. Cells from the dorsal base of a leaf, X 310. 6. Cells from 
the ventral base of a leaf, X 310. 7. Cells from the basal portion of a leaf, 
X 310. 8. Underleaf, X 30. 9. Cells from a lobe of an underleaf, X 310. 
10. Female bracts of an outer series, X 30. 11. Female bracts of the inter- 
mediate series, X 30. 12. Female bract of the innermost series, X 30. 13. One 
of the laciniae of a bract of this series, X 90. 14. Portion of the mouth of 
the perianth (immature), X 90. 15. Portion of a plant, ventral view, X 15. 
16. A cell from the apical portion of a leaf, X 400. 17. Underleaf, X 30. 
18. Portion of a plant, ventral view, X 15. 19. A cell from the apical portion 
of a leaf, X 400. 20. Underleaf, X 30. Nos. 1-14 drawn from a portion of 
the type, from Guadeloupe; 15-17 from plants of forma paupercula, from 
Brazil; 18-20 from a portion of the type of M. Martianum, from Brazil. 



M. Fulford 62 Bazzania 

long, 0.8 mm. - 1 mm. broad at the base, narrowed a little to the 
transversely truncate, tridentate apex: the dorsal margin arched 
from a rounded base, covering one-half the stem, the ventral 
margin a little concave, the base rarely dilated, the apex irregularly 
three-toothed, the teeth mostly large, acute, four to six cells long 
and broad, the sinuses deep, acute to lunulate, the margins entire ; 
leaf cells thin-walled, the trigones large, with bulging sides, often 
coalesced, the cell lumina angular-rounded to stellate, the cuticle 
slightly verruculose; cells of the apical region 22/x X 22/i, of the 
interior larger, and of the base 40/,t - 48/A X 22/z, a vitta not dif- 
ferentiated : underleaves approximate to imbricated, attached in 
a straight line, subquadrate, broader than the stem, mostly 0.56 
mm. long X 0.7 mm. broad, the lateral margins deeply lobed, the 
apex distinctly four-lobed, the lobes broad, rounded, mostly 3 to 8 
cells long, 6 to 8 cells broad, a slime papilla usually present at the 
apex of each lobe, the cells as in the leaf : leaves of the flagellif orm 
branches ovate, scale-like: male branches several on a stem, the 
bracts and bracteoles round-quadrate, bi-trifid, antheridia occurring 
singly: female branches occasional, solitary, one to several on a 
stem, the bracts and bracteoles similar, the outer series ovate, the 
apex shortly bi-trifid ; the intermediate series ovate, longer, to one- 
fourth divided into two or three long, often serrulate teeth, the 
lateral margins serrulate to dentate; the innermost series long 
ovate, the apex divided into usually three, long, serrulate laciniae ; 
the perianths mostly 5 mm. long, the mouth ciliate to laciniate. 

HABITAT : In mats on logs and tree bases. 

The distinguishing characteristics of the species are its large 
size, the olive-green to brown color, the tridentate leaves with con- 
spicuous trigones and angular-rounded to stellate cell lumina, and 
the deeply four-lobed, subquadrate underleaves. (FiG. 19, 
nos. 1-20). 

The forma paupercula collected by G. A. LINDBERG at Caldas, 
Brazil is much smaller, and shows evidence of having grown under 
unfavorable habitat conditions. The trigones are not so large but 
are, nevertheless, of the same form. The underleaves are smaller 
and while some of them are nearly entire with only a suggestion 
of lobing at the apex, others on the same plant are deeply four- 
lobed and identical with the underleaves on the large plants (see 
FIG. 19, nos. 15-17). 

Mastigol)i*yum Martianum Gottsche, also collected by G. A. 
LINDBERG at Caldas, Brazil is identical with the form paupercula 
mentioned above (compare FIG. 19, nos. 15-17 with nos. 18-20), 
and is therefore also to be considered a depauperate form of B. 
quadricrenata. 

DISTRIBUTION: Guadeloupe: without locality, 1'Herminier, the type 
(G). Jamaica : John Crow Peak, Maxon 1240 (Y) ; Morce's Gap, Evans 
465 (Y). Brazil : Caldas, (forma paupercula) , G. A. Lindberg (G); 
also type of M. Martianum (NY). Venezuela: without locality, 
Korthals (G). 

REFERENCES: Stephani (1908, 508; 1909, 525; /cones, Mastigobryum nos. 
348, 393); Bescherelle (1893, 186). 



Bazzania 63 Grandistipulae 

15. Bazzania aurescens Spruce, Trans. & Proc. Bot. Soc. [Edin- 
burgh] 15: 374. 1885. 

Mastigobryum aurescens Stephani, Spec. Hep. 3: 507. 1908. 
Bazzania Hookeri var. Spruce ms., Hepat. Sprue. 

Plants medium to large, greenish brown, becoming dark yellow- 
brown in the older portions : stems slender, 6 cm. or more in length, 
with leaves to 3.5 mm. broad, prostrate to ascending; lateral 
branches infrequent, diverging at a wide angle; ilagelliform 
branches frequent, long ; rhizoids colorless, present on some leaves 
of the flagellif orm branches : the line of leaf insertion curved in its 
upper part; leaves approximate to subimbricated, plane, becoming 
deflexed when dry, unsymmetrically narrow-ovate to oblong, spread- 
ing, 1.5 mm. - 2 mm. long, 0.7 mm. - 0.8 mm. broad at the base, nar- 
rowed a little to the transversely truncate, tridentate apex; the 
dorsal margin arched from a curved base, covering one-half the 
stem, the ventral margin straight to a little concave, the base 
scarcely dilated, the apex three-toothed, the teeth irregular, three to 
six cells long and broad, acute to acuminate, the sinuses lunulate, the 
margins usually crenulate; leaf cells thin- walled, the cell lumina 
angular-rounded to stellate, the trigones large, with convex sides, 
often becoming coalesced, the cuticle faintly verruculose ; cells of the 
apical region 20/t - 24//, in diameter, of the median portion larger, 
and of the basal portion to 40/A X 20/t, a vitta not differentiated : 
underleaves imbricated, subquadrate in outline, broader than the 
stem, the line of attachment straight, 0.6 mm. - 0.7 mm. long and 
broad, the lateral margins a little convex from a straight base, 
repand, the apex irregularly incised with unequal teeth and lobes, 
the cell pattern as in the leaf: leaves of the flagelliform branches, 
scale-like, ovate, the apex acute to shortly bifid: female branches 
occasional, solitary, one to several on a stem, the bracts and 
bracteoles similar, ovate ; the outermost series short, the apex bifid, 
the intermediate series larger, divided to one-fourth into three 
long narrow laciniae, the innermost series long, one-sixth to one- 
fourth divided into three ciliate laciniae, the lateral margins den- 
tate to short ciliate: perianth (immature) mouth fringed with long 
cilia: male branches not seen. 

HABITAT : On shaded rocky slopes on Mount Guayrapurina, also 
on logs at higher altitudes in Jamaica. 

The distinguishing characteristics of the species are its greenish 
brown color ; the oblong to ovate leaves with transversely truncate, 
equally tridentate apices which have irregular, mostly crenate 
margins, and cells with thin walls and very large trigones ; and the 
subquadrate underleaves, repand on the lateral margins and incised 
to form irregular teeth or lobes at the apex. (FlG. 20, nos. 1-11). 

The plants of the type material are quite large and the yellow- 
brown pigmentation is strongly developed. The teeth of the leaves 
are long, usually acuminate, and the margins are conspicuously 
crenulate as is seen in FIG. 20, nos. 1, 3 and 7. This form of margin 
is present to some degree in all leaves but in the less well developed 
forms it is less obvious. The large cells with their very large, 




FIG. 20. Bazzania aurescens Spruce. 1. Portion of a plant, ventral 
view, X 15. 2. Portion of a leaf and stem, dorsal view, X 30. 3. A tooth 
of a leaf, X 310. 4. A cell from the apical portion of a leaf, X 400. 5. Under- 
leaves, X 30. 6. A portion of the apical margin of an underleaf, X 310. 
7. Apices of leaves, X 30. 8. Female bract of the outermost series, X 30. 
9. Female bract of the innermost series, X 30. 10. One of the laciniae of a 
bract of this series, X 310. 11. Portion of the mouth of a perianth, X 100. 
Nos. 1-6 drawn from a portion of the type material, from Peru; 7-11 from 
plants of the type of B. Hookeri var. Spruce, also from Peru. 



Bazzania 65 Grandistipulae 

rounded trigones, are another conspicuous feature of the leaves. The 
cell walls are thin but usually this is not obvious because of the de- 
position of abundant secondary thickenings, so that only small, thin- 
walled pits remain (FiG. 20, nos. 3 and 4). The cell lumina are 
stellate. 

The underleaves, FIG. 20, nos. 1, 5 and 6, are large. They are 
subquadrate in outline and are variously lobed and often irregularly 
toothed, with broad, blunt teeth along the margins. The cells are 
similar to those of the leaf. 

Plants similar to those described above were distributed in the 
Hepaticae Spruceana as B. Hookeri var. Some of these plants 
have female branches in various stages of development. The 
laciniae of the intermediate and innermost series of bracts are 
long, and are short ciliate along the margins (see FIG. 20, nos. 
8-10). The perianth mouth is fringed with numerous long cilia 
as is shown in FIG. 20, no. 11. 

Plants from Jamaica are much smaller than those from South 
America. The stems with leaves average only 2.5 mm. - 3 mm. 
broad and are deeply pigmented with brown. The teeth of the 
leaves are not nearly so long, but they are sharp, and many of 
them have more or less crenulate margins. The female bracts 
and the perianth mouths are like those of the plants from Peru. 
Many of the underleaves were not nearly so lobed and dentate as 
those shown in FIG. 20, nos. 1 and 5, but most of them showed 
some indication of lobing along the apical margins with two or 
more teeth at the apex. In this condition they are very similar to 
the underleaves of B. Glaziovii from Brazil. However, this latter 
species usually is more robust than even the plants from Peru and 
the teeth of the leaves are rather short and broad, with the margins 
not crenulate. Female branches and perianths have not been found 
in B. Glaziovii so that no comparisons of these characters can be 
made at this time. 

Poorly developed plants may be confused with depauperate 
forms of B. quadricrenata. However, the larger cells and very 
large trigones of the leaves of B. aurescens should aid in the sepa- 
ration of the two species. If female branches are present the 
mouth of the perianth will furnish additional characteristics. 

DISTRIBUTION: Jamaica: Morce's Gap, Underwood 622 (NY); John 
Crow Peak, Underwood 678, 2364 (NY). Peru: Mt. Guayrapurina, 
Spruce, Hepat. Sprue., the type (H) ; the same, as B. Hookeri var. (NY). 
Venezuela : without locality, Fendler (as M. scutigerum), poor (H). 

REFERENCES: Stephani (Icones, Mastigobryum no. 343). 

16. Bazzania Glaziovii (Gottsche) comb. nov. 

Mastigobryum Glaziovii Gottsche in Stephani, Hedwigia 25: 8. pi. 4. fig. 1-2. 
1886. 

Plants medium to large, dark green, becoming pigmented with 
brown in the older portions: stems stout, to 5 cm. or more in 




FIG. 21. Bazzania Glaziovii (Gottsche) Fulford. 1. Portion of a 
plant, ventral view, X 15. 2. A leaf, X 15. 3. A tooth of a leaf, X 310. 
4. A cell from the apical portion of a leaf, X 400. 5. Underleaf, X 15. 6. A 
portion of the apical margin of an underleaf, X 310. Drawn from the type 
material. 



Bazzania 67 Grandistipulae 

length, with leaves to 4 cm. broad, prostrate to suberect; lateral 
branches frequent, 1 cm. or more apart, diverging at a wide angle ; 
flagelliform branches frequent; rhizoids colorless, present on the 
bases of the leaves of the flagelliform branches; the line of leaf 
insertion curved in the upper part; leaves imbricated, plane to 
deflexed, unsymmetrically ovate, spreading, 1.8 mm. - 2.3 mm. 
long, 1 mm. broad at the base, narrowed to the more or less 
obliquely truncate, tridentate apex; the dorsal margin strongly 
arched from a cordate base, extending across the stem and some- 
times beyond, the ventral margin straight to concave, the base 
scarcely dilated, the apex irregularly three-toothed, the teeth broad, 
three to five cells high, four to seven cells broad at the base, the 
sinuses lunulate, the margins entire ; leaf cells thin- walled, the cell 
lumina angular-rounded, the trigones large, distinct, often becom- 
ing coalesced, the cuticle smooth to faintly verruculose; cells of 
the apical portion and dorsal base mostly 20u - 24/i in diameter, 
those of the median portion larger, those or the base 40/1 - 48/i 
X 24/i, a vitta not differentiated: underleaves imbricated, sub- 
quadrate in outline, attached in a straight line, broader than the 
stem, 0.7 mm, - 0.85 mm. long, 0.6 mm. - 0.7 mm. wide, the lateral 
margins somewhat convex from a curved base, often lobed, the 
apex usually with a short, incurved tooth at either end, undulate 
to lobed between, the cells as in the apex of the leaf: leaves of the 
flagelliform branches scale-like, ovate : sexual branches not seen. 

HABITAT : Not given. 

The distinguishing characteristics of the species are its dark 
green color and large size ; the large, obliquely truncate leaves with 
large, thin-walled cells and conspicuous trigones; and the large, 
subquadrate underleaves with entire or lobed lateral margins and 
truncate, several lobed apices which have an incurved tooth at 
either end. (FiG. 21, nos. 1-6). 

The species is distinguished from well developed forms of B. 
qiuidricrenata because of the deeply f our-lobed apices of the under- 
leaves of the latter. Poorly developed plants of the two species are 
very similar. The leaves, with their shorter, broader teeth with 
smaller cells and smaller trigones will separate it from B. 
aurescens. 

The underleaves offer the most accurate guide in the recogni- 
tion of the species. They are large, broader than the stem and 
usually imbricated. The lateral margins are undulate to irregu- 
larly lobed. The apex is truncate and usually has a short, incurved 
tooth on each end, with several lobes between as is seen in FIG. 21, 
nos. 1, 5 and 6. The teeth of the leaves are always short and broad 
and separated by shallow sinuses (see FIG. 21, nos. 1, 2 and 3). 

DISTRIBUTION: Brazil: Rio de Janeiro, Glaziou, the type (H). 
British Guiana: Turkeit, Lutz (NY). 

REFERENCES: Stephani (1909, 528; /cones, Mastigobryum no. 386). 



M. Fulford 68 Bazzania 

17. Bazzania Breuteliana (Lindenb. & Gottsche) Trevis. Mem. 1st. 

Lomb. 13: 414. 1877. 

Mastigobryum Breutelianum Lindenberg & Gottsche, in G. L. & N., Syn. Hep. 

226. 1845. 

Bazzania vincentina var. subrectifolia Spruce ms. p.p. 
Mastigobryum portoricense Hampe & Gottsche, Linnaea 25: 348. 1852. 
Mastigobryum Cuervi Gottsche, Ann. Sci. Nat. Bot. Ser. V. 1 : 141. 1864. 
Mastigobryum sylvaticum Gottsche, Husnot, Hep. Exsicc. (nomen nudum). 

Not M. sylvaticum Stephani. 
Bazzania chimb or azensia Spruce, Trans. & Proc. Bot. Soc. [Edinburgh] 15: 

376. 1885. 

Mastigobryum chimborazense Stephani, Spec. Hep. 3: 527. 1909. 
Btizzania viridissima Spruce, op. cit. p. 375. 
Mastigobryum viridissimum Stephani, op. cit. p. 522. 
Mastigobryum Uleanum Stephani, op. cit. p. 529. 
Bazzania portoricensis var. pycnodictyon Herzog, Rev. Bryol. et Lichen. 11: 

19. 1938. 
Mastigobryum Hariotii Stephani, in Herzog, Biblioth. Bot. 87: 224. Fig. 164, 

d-e. 1916. 

Plants large, in deep tufts or mats, olive-green to brownish 
green, pigmented with brown : stems to 10 cm. or more in length, 
with leaves to 4.5 mm. broad, prostrate to suberect; stem cells in 
longitudinal section to 0.18 mm. long, the cortical shorter, both 
averaging 20/x in diameter, the vertical walls uniformly thickened 
and containing frequent pits, the end walls thin : lateral branches 
frequent, mostly more than 5 mm. apart, diverging at a wide angle ; 
flagelliform branches frequent: rhizoids colorless, rare, on the 
bases of the leaves of the flagelliform branches: the line of leaf 
insertion curved in its upper part, the dorsal margin recurved 
forming a short hook; leaves distant to imbricated, plane to de- 
flexed, unsymmetrically ovate, oblong on robust plants, spreading, 
often becoming a little falcate, 1.5 mm. to 2.6 mm. long, 0.8 
mm. - 1 mm. broad at the base, narrowed a little to the trans- 
versely truncate, unequally tridentate apex; the dorsal margin 
arched from a strongly rounded base, extending the width of the 
stem and often beyond, the ventral margin straight to a little con- 
cave, the base scarcely dilated, the apex irregularly three-toothed, 
the teeth large, acute, two to eight cells long and broad, occasion- 
ally obscure, the sinuses deep, acute to lunulate, the margins entire ; 
leaf cells thin-walled, the cell lumina angular-rounded, the trigonea 
small, conspicuous, the cuticle verruculose ; the cells tending to be 
in rows, those of the apical region and dorsal base 20/* X 20/x, of 
the interior larger, and of the base 36ft - 54ft X 18/x, a vitta not 
differentiated: underleaves distant to imbricated, subquadrate to 
elongate in outline, attached in a straight line, much broader than 
the stem, 0.8 mm. - 1 mm. long, 0.65 mm. - 1 mm. wide, the lateral 
margins convex, entire, repand, the base cordate, the apex usually 
two- to four-lobed, the lobes broad, rounded, the cells as in the 
leaf: leaves of the flagelliform branches spreading, small, ovate, 
the apex acute to shortly bifid : female branches occasional, solitary, 
one to several on a stem, the bracts and bracteoles similar, the 
outermost series oblong, bifid, the intermediate series ovate, the 
apex divided into three or four short, serrulate laciniae, the lateral 
margins serrulate to dentate ; the innermost series ovate, one-third 




FIG. 22. Bazzania Breuteliana (Lindenb. & Gottsche) Trevis. 1. Por- 
tion of a leaf and stem, dorsal view, X 30. 2. Portion of a plant, ventral view, 
X 12. 3. A leaf. X 30, 4. Apices of leaves, X 30. 5. A cell from the apical 
portion of a leaf, X 350. 6. Cells of an apical tooth, X 260, 7. Cells from 
the dorsal base, X 260. 8. Cells from the basal portion, X 260. 9. Under- 
leaves, X 30. 10. Cells from the apical portion of an underleaf, X 260. 
11. Portion of a cross-section of a stem, X 260. Nos. 1-11 drawn from the 
type material. 



M. Fulford 70 Bazzania 

to one-fourth divided into three or four long, serrulate laciniae, 
the margins serrulate to short ciliate or laciniate : perianth mouth 
short laciniate : male branches not seen. 

HABITAT : On logs, tree bases and branches in woods. 

The distinguishing characteristics of the species are its large 
size, and olive-green color faintly tinged with brown; the large, 
spreading to slightly falcate leaves with irregularly tridentate 
apices, angular-rounded cell lumina, thin cell walls, and small, dis- 
tinct trigones; and the quadrate to elongate, faintly two- to four- 
lobed underleaves, cordate at the base, with mostly repand lateral 
margins which are straight or a little convex. (FIGS. 22, nos. 1-11 ; 

23, nos. 1-14; 24, nos. 1-11). 

The plants exhibit the usual variations associated with differ- 
ences in habitat. The stems are usually long, branched or un- 
branched, with the branches mostly from 0.5 mm. to 1.5 mm. 
apart. They are olive-green with a brown pigmentation which 
becomes more conspicuous with age. The leaves are distant to 
imbricated, unsymmetrically ovate (on the type material, FIG. 22, 
no. 3), to elongate on the robust plants (FlGS. 23, nos, 1, 7 and 9; 

24, 3) and are usually spreading, even when dry. The teeth vary 
in size and shape even on a single stem. The cells always have 
thin walls and small, distinct trigones. In plants of exposed situa- 
tions secondary thickenings are laid down to the extent that some 
of the trigones become coalesced. 

The underleaves show a wide degree of variation. They may 
be distant on the unbranched elongate stems, but are usually im- 
bricated. The bases are rounded from a straight line of attach- 
ment. On well developed plants definite auricles are formed but 
usually the bases are only rounded, and in a few instances are 
nearly straight. However, at least a few underleaves of a plant 
will show the well-rounded condition. The lateral margins are 
mostly a little convex, entire to undulate. On robust plants they 
are plane or only a little repand, but on the smaller, little branched 
plants they become strongly repand. The apex is characteristically 
broad and three- or four-lobed. These lobes are short and broad and 
the sinuses are lunulate. Some underleaves on a plant always show 



FIG. 23. Bazzania Breuteliana (Lindenb. & Gottsche) Trevis. 1. Por- 
tion of a plant, ventral view, X 15. 2. Portion of a leaf and stem, dorsal 
view, X 30. 3. A cell from the apical portion of a leaf, X 400. 4. Portion 
of a plant, ventral view, X 10. 5. A cell from the apical portion of a leaf, 
X 400. 6. Underleaf, X 30. 7. Portion of a plant, ventral view, X 15. 8. A 
cell from the apical portion of a leaf, X 400. 9. Portion of a plant, ventral 
view, X 15. 10. A cell from the apical portion of a leaf, X 400. 11. Portion 
of a plant, ventral view, X 15. 12. Cells from the apical portion of a leaf, 
X 400. 13. Portion of a stem, ventral view, X 15. 14. A cell from the apical 
portion of a leaf, X 400. Nos. 1-3 drawn from a portion of the type of 
Af. port&ricense, from Puerto Rico; 4-6 from a portion of the type of Af. 
CuervL from Colombia; 7-8 from a portion of the type of B. chimborazensis, 
from Ecuador; 9-10 from a portion of the type of B. viridissima, from Peru; 
11-12 from a portion of the type of Af. Uleanum, from Peru; 13-14 from a 
portion of the type of B. portoricensis var. pycnodictyon, from Costa Rica. 




FIG. 24. Bazzania Breuteliana (Lindenb. & Gottsche) Trevis. 1. Por- 
tion of stem, ventral view, X 12. 2. Dorsal side of stem showing leaf attach- 
ment, X 30. 3. Leaf, X 30. 4. Underleaf, X 30. 5. Female bract of outermost 
series, X 30. 6-7. Female bracts of the intermediate series, X 30. 8. A tooth 
of a bract of an intermediate series, X 100. 9. Female bracts of the inner- 
most series, X 30. 10. Portion of one of the laciniae of a bract of this series, 
X 100. 11. Portion of the mouth of a perianth (immature), X 100. Nos. 
1-11 drawn from plants collected by WEBSTER in Jamaica. 



Bazzania 73 Grandistipulae 

the characteristic lobed condition (FiG. 22, no. 9; FiG. 23, nos. 1, 
4, 7, 9 and 11), of the apex. Female bracts and perianths were 
abundant on plants collected in Jamaica (see FIG. 24, nos. 5-11). 
When present, these will furnish additional diagnostic characters. 

The plants of the type material (see FIG. 22, nos. 1-11), from 
St. Kitts are robust but the leaves are somewhat shorter than those 
of the other collections. They are very broad at the base, so that 
they are unsymmetrically ovate rather than elongate. Most of the 
teeth of the broad leaf apices are short and broad and separated 
by narrow sinuses. However, some of the leaves on a stem have 
much longer or larger teeth so that the short teeth cannot be used 
as a specific characteristic. The underleaves are large, rounded at 
the bases and uniformly truncate and three- or four-lobed or 
undulate at the apices. 

The plants of the type collection of M. portoricense (FiG. 23, 
nos. 1-3), from Puerto Rico, are a little larger and a little more 
deeply pigmented than those of M. Breutelianum. Most of the 
leaves are elongate. The underleaves are perhaps a little longer 
than those described above, but the apices are lobed in the 
characteristic manner. 

M. Cuervi (FiG. 23, nos. 4-6), from Colombia, is a relatively 
small form with short, ovate leaves. The teeth are irregularly 
developed, some very short and broad, others very long and acute. 
The underleaves are distant but have the same characteristic apices. 
The cell walls are somewhat thicker and the trigones a little larger 
than those of the leaves of M. Breutelianum or M. portoricense but 
the general cell pattern of the three is identical. 

Plants of B. chimborazensis (FiG. 23, nos. 7-8), from Ecuador, 
are very large as is seen by a comparison of the habit sketches of 
FIGS. 22 and 23, no. 7. The leaves are elongate and very sharply 
toothed. The contrast between the size of these teeth and those of 
M. Breutelianum is so striking that at first glance one is apt to 
consider the two as distinct species. However, an examination of 
many plants from the West Indies and South America has revealed 
an overlapping series of gradations in the size of the teeth on 
individual plants, and M. Breutelianum with most of the teeth very 
short and broad is at one end of the series, while B. chimborazensis 
with most of the teeth large, is at the other. Some of the leaves of 
these plants have shorter teeth. The underleaves are very large, 
but typical for the species. 

The plants of M. Hariotii from Bolivia also have sharply 
toothed leaves. The stems are not nearly so large as those of B. 
chimborazensis and the leaves are shorter and broader at the bases. 
The underleaves are distant. 

B. viridissima (FiG. 23, nos. 9-10), from Peru, is anotfier 
example of a very large form of the species. The teeth of the 
leaves are very large, often longer than those of B. chimborazensis. 
The underleaves are typical. 



M. Fulford 74 Bazzania 

The plants of M. Uleanum (FiG. 23, nos. 11-12), also from Peru, 
are more similar to M. Breutelianum than to the other South 
American plants described above. The leaves, while elongate, are 
for the most part tridentate with short, broad teeth. The under- 
leaves are mostly as broad as long, cordate at the base, undulate 
along the margins and three- or four-lobed at the apices. The 
presence of this form in Peru further substantiates the very close 
relationship of the various plants described above and besides, 
seems to indicate that the plants with very large teeth, B. chim- 
borazensis and B. viridissima do not necessarily make up a 
geographic variety. Many of the representatives of the species of 
Bazzania which SPRUCE collected in the Amazon country of South 
America have the leaves more deeply toothed than plants of those 
same species collected elsewhere in tropical America. 

The plants of B. portoricensis var. pycnodictyon (FiG. 23, nos. 
13-14), from Costa Rica, differ only in that the cuticle is strongly 
verruculose. But since the cuticle of the plants of M. Breutelianum 
is slightly verruculose this characteristic seems not to be of suffi- 
cient importance to warrant a varietal name. The female bracts 
are identical with those of the species. 

The species can readily be separated from the preceding species 
because of its large size, and large, subquadrate to elongate under- 
leaves with entire or faintly undulate lateral margins, rounded or 
cordate at the bases from a straight line of insertion, and the trun- 
cate, three- or four-lobed apices. Small forms might sometimes 
be confused with B. acuminata, a description of which follows. 

DISTRIBUTION: Cuba: Oricnte, Shafer 9116 (Y, NY). Dominica: 
without locality, Eggers, cited by Stephani (1888b, 300); Morne Micotrin, 
Roseau Valley, Morne Diablotin, Elliott, cited by Spruce (1895, 356). 
Dominican Republic: Pacificador, Abboy 2061a, 2063, 2126 (Y). 
Guadeloupe : Morne Hirondelle, Duss 282 (NY); without locality, Hus- 
not, the type of M. sylvaticum Gottsche (H) ; without locality or collector's 
name (H); without locality, 1'Herminier, Soufriere, Duss, cited by Stephani 
(1903, 22). Jamaica: without locality, Webster (NY); Sir John's 
Peak, Harris 11, 135a (NY); Morce's Gap, Evans 41, 44, 390 (Y). 
Martinique: Mt. Pelee, Duss 126, 127, 347 (NY) ; without locality, Hahn, 
cited by Bescherelle (1893, 186). Puerto Rico: without locality, 
Schwanecke, the type of M. portoricense (NY). El Yunque, Evans 11, 66, 
132 (NY); Sierra Naguabo, Shafer, a, 3306, 3455, 3729 (NY); same local- 
ity, E. G. Britton and Hess 2291 (NY); El Yunque, P. R. College of Agri. 
2792 (NY); Mt. Britton, Jones 10976 (H); Alto de la Bandera, Britton and 
Marble 2175 (NY); Luquillo Mountains, E. G. Britton 7753 (NY) ; Can6vanas, 
Pagan 291 (NY); Adjuntas, Sintenis 92, as B. Wrightii (NY); without 
locality, Sintenis 28, 87, cited by Stephani (1888a, 279). St. Kitts : Mt. 
Misery, Britton & Cowell 783 (NY); Mt. Misery, Breutel, the type. 
Mexico : Oaxaca, Liebmann 255 a & b, plants mixed with M. taleanum 
(G, H). Costa Rica : San Marcos de Data, Tonduz 11618 (NY); La 
Estrella 39106, 39117, 39403; El Muneco, Standley 51338, 50887, the type of 
B. portoricensis var. pycnodictyon (W). Bolivia: Casapi, Mathews 
(H, NY); Comarapa, Herzog, the type of M. Hariotii (L). Brazil : 
Mangos, Ulev the type of M. Uleanum (H); without locality, Sells as M. 
Martianum (H). Colombia: Bogota, Cuervo, the type of M. Cuervi 
(B) ; without locality, Wallace (NY). British Guiana: Mt. Roraima, 



Bazzania 75 Grandistipulae 

McConnell & Quelch 545, cited by Stephani (1901-05, 98). Ecuador: 
Tunguragua, Spruce, Hepat. Sprue, as B. vincentina, var. subrectifolia p.p. 
(NY); Mt. Chimborazo, Spruce, the type of B. chimborazensis (NY). 
Peru : Campana, Spruce, the type of B. viridissima (H). 

REFERENCES: Herzog (1938, 19); Lindenberg & Gottsche (1851, 75); 
Stephani (1888a, 279; 1888b, 300; 1901-05, 98; 1903, 22; 1908, 470; 1909, 519, 
522, 526; Icones Mastigobryum nos. 157, 164, 378> 382, 397a, 6, 403, 407). 

18. Bazzania acuminata (Lindenb. & Gottsche) Trevis. Mem. 1st. 

Lomb. 13: 414. 1877. 

Mastigobryum acuminatum Lindenberg & Gottsche, in G. L. & N., Syn. Hep. 

719. 1847. 

Mastigobryum orizabense Gottsche, Mex. Lebeim 130. 1863. 
Mastigobryum Mullerianum Gottsche, op. cit. p. 129. 

Plants medium size, olive-green, deeply pigmented with brown : 
stems to 5 cm. long, with leaves to 2.5 mm. or 3 mm. broad; 
lateral branches frequent, diverging at an acute angle ; flagelliform 
branches frequent, long : rhizoids colorless, abundant on the leaves 
of the flagelliform branches: the line of leaf insertion curved in 
its upper half, the dorsal extremity curved downward, forming a 
short hook; leaves distant to imbricated, strongly deflexed when 
dry, unsymmetrically and broadly ovate, ascendent, 1.5 mm. - 2 mm. 
long X 0.8 mm. - 1 mm. broad at the base, narrowed to the mostly 
obliquely truncate, unequally tridentate apex; the dorsal margin 
strongly arched from a cordate base, extending across the axis and 
somewhat beyond, the ventral margin nearly straight, the base 
scarcely dilated, the apex irregularly two- or three-toothed, the 
teeth acute, two to five cells long and broad, the sinuses mostly 
shallow, lunulate, the margins entire; leaf cells thin-walled, the 
trigones conspicuous, with bulging sides, sometimes coalesced, the 
cell lumina angular-rounded, the cuticle verruculose; cells of the 
apical region 18/x X 18/t, those of the interior larger, and those of 
the base 36/t - 46/A X 18/x, a vitta not differentiated: underleases 
distant to approximate, orbicular, attached in a straight line, 
broader than the stem, 0.5 mm. - 0.56 mm. long X 0.45 mm. - 0.52 
mm. broad, the lateral margins entire, strongly recurved, the apex 
entire, emarginate, sometimes with several undulations, the cells 
as in the leaf : leaves of the flagelliform branches ovate, scale-like, 
spreading: sexual branches not seen. 

HABITAT : Not given. 

The distinguishing characteristics of the species are its medium 
size, the brown pigmentation, the unsymmetrical, broadly ovate 
leaves with strongly cordate dorsal bases and hook-formed lines of 
attachment, and the orbicular, emarginate underleaves with 
strongly recurved lateral margins. (FiG. 25, nos. 1-11). 

The type collection, from Mexico, in the Herb. Boissier contains 
two fragments of stems each less than 1 cm. in length. The 
characteristic hooked line of insertion of the leaves, the leaf shape, 
the trigones, and the orbicular, emarginate underleaves with re- 
curved lateral margins are distinctive (see FIG. 25, nos. 1-6). 




/A - ' ' \ ' , \ 

6 \V k' v I' ' 



FIG. 25. Bazzania acuminata (Lindenb. & Gottsche) Trevis. 1. Por- 
tion of stem, ventral view, X 30. 2. Dorsal side of stem to show leaf attach- 
ment, X 30. 3. Leaf apices, X 30. 4. A cell from the apical portion of a leaf, 
X 400. 5. Underleaves, X 30. 6. Cells from the apical portion of an under- 
leaf, X 310. 7. Portion of a plant, ventral view, X 15. 8. A cell from the 
apical portion of a leaf, X 400. 9. Underleaf, X 30. 10. Portion of a plant, 
ventral view, X 15. 11. A cell from the apical portion of a leaf, X 400. Nos. 
1-6 drawn from a portion of the type, from Mexico; f-9 from the type of M. 
orizabense, from Mexico; 10-11 from the type of M. Miillerianum, from Mexico. 



Bazzania 77 Grandistipulae 

The type collection of M. orizabense (FiG. 25, nos. 7-9), also 
from Mexico contains several stems, all with the same character- 
istics as are present in M. acuminatum. These stems are in a much 
better state of preservation. 

M. Mullerianum (FiG. 25, nos. 10-11), from Mexico, is a little 
less pigmented with brown than M. orizabense, and some of the 
leaves have more conspicuous teeth, but the variation is so great, 
even on one stem that this characteristic is not of sufficient stability 
to separate M . Mullerianum as a distinct species. 

The species is most similar to small plants of B. Breuteliana. 
However, the emarginate underleaves with strongly recurved lat- 
eral margins will usually serve to distinguish B. acuminata from 
all other species. In addition to this, the underleaves of B. 
Breuteliana, even though the lateral margins may be slightly re- 
curved near the base, are always subquadrate to elongate in outline, 
rather than orbicular, and the truncate apices are three- or four- 
lobed rather than emarginate. 

DISTRIBUTION: Mexico : Oaxaca, Liebmann 176b, the type (G) ; Vera 
Cruz, Orizaba, Muller 159, the type of M. orizabense (G) ; Orizaba, Miiller 
2361, the type of M. Mullerianum (NY). Venezuela: Caracas, 
Burchel (NY). 

REFERENCES: Lindenberg- & Gottsche (1851, 69); Gottsche (1863, 131); 
Stephani (1909, 517, 518;, Ic<mes, Mastiffobryum nos. 370, 394a> b, 395). 

19. Bazzania diversicuspis Spruce, Trans. & Proc. Bot. Soc. [Edin- 
burgh] 15: 373. 1885. 

Mastigobryum diversicuspe Stephani, Spec. Hep. 3: 432. 1908. 

Plants delicate, in depressed mats or scattered among other 
bryophytes, dull green to light brown: stems slender, to 4 cm. or 
5 cm. long, with leaves to 3 mm. broad, prostrate; stem cells in 
longitudinal section elongate, to 0.17 mm. in length, the cortical 
shorter, both averaging 20/x in diameter, the vertical walls uni- 
formly thickened and containing frequent pits, the end walls thin : 
lateral branches 1 cm. or more apart, diverging at a wide angle; 
flagelliform branches frequent, long: rhizoids colorless, present 
on the leaves of the flagelliform branches : the leaf insertion little 
curved in the upper part; leaves spreading, approximate to im- 
bricated, straight to ascendent, unsymmetrically ovate, to 1 mm. 
long, 0.48 mm. broad at the base, narrowed somewhat to the acute, 
rounded or transversely truncate apex, the dorsal margin convex 
from a slightly rounded base which covers one-half the stem, the 
ventral margin straight to slightly concave, the base scarcely 
dilated, the apex mostly broad, acute, rounded or obscurely bi- or 
tridentate ; teeth when present, acute, three to eight cells broad at 
the base, two to four cells high, the sinuses lunulate, the margins 
entire ; leaf cells more or less quadrate in outline, thin-walled, the 
trigones small, conspicuous, seldom becoming coalesced, the cuticle 
verruculose; cells of the apical region and dorsal base mostly 
17/A X 17/i, those of the median portion a little larger, those of the 
base 27/A - 36ju X 18/t, not forming a vitta : underleaves distant to 




FIG. 26. Bazzania diversicuspis Spruce. 1. Portion of a stem and 
leaf, dorsal view, X 30. 2. Portion of a plant, ventral view, X 30. 3. A 
leaf, X 30. 4. A cell from the apical portion of a leaf. X 350. 5. Cells from 
an apical tooth of a leaf, X 260. 6. Cells from the dorsal margin near the 
base, X 260. 7. Underleaves, X 30. 8. Cells from the apical portion of an 
underleaf, X 260. Nos. 1-8 drawn from a portion of the type collection. 



Bazzania 79 Grandistipulae 

approximate, attached in a straight line, subquadrate in outline, a 
little broader than the stem, 0.28 mm. - 0.36 mm. long and broad, 
the lateral margins a little convex from a straight base, the apex 
straight, undulate, two- to four-lobed or occasionally with one or 
two short, sharp teeth, cells similar to those of the apical portion 
of the leaf: leaves of the flagelliform branches scale-like, acute, 
0.9 mm. long : male and female branches not seen. 

HABITAT : In depressed mats among ferns. 

The distinguishing characteristics of the species are its dull 
green to brownish color, the small to medium size, the prostrate 
habit ; the plane, mostly obscurely bi- or tridentate leaves with thin 
cell walls; and the small, quadrate, entire or faintly lobed under- 
leaves. (FiG. 26, nos. 1-8). 

The obscurely tridentate, acute or bidentate leaves and the 
small, distant to approximate underleaves will readily separate B. 
diversicuspis from all of the preceding species. 

DISTRIBUTION: Trinidad : without locality or collector's name (NY). 
Brazil : Tauau near Para, Spruce, Hepat. Sprue., the type (NY). 

REFERENCES: Stephani (1886, 244; 1908, 432; Icones, Mastigobryum no. 8). 

20. Bazzania tricuspidata (Steph.) comb, nov* 
Mastigobryum tricusjndatum Stephani, Spec. Hep. 3: 448. 1908. 

Plants small, golden brown, becoming darker in the older por- 
tions : stems slender, 3 cm. or more in length, with leaves to 2 mm. 
broad ; lateral branches mostly more than 1 cm. apart, diverging at 
a wide angle ; flagelliform branches occasional ; rhizoids numerous 
from the bases of the leaves of flagelliform branches : the line of 
leaf insertion little curved in its upper part ; leaves distant, strongly 
deflexed, unsymmetrically ovate, ascendent, mostly 1 mm. long', 
0.5 mm. broad at the base, narrowed a little to the broad, trans- 
versely truncate, tridentate apex; the dorsal margin arched from 
a straight base, covering one-half the stem, the ventral margin 
mostly straight, the base scarcely dilated, the apex more or less 
equally three-toothed, the teeth three to six cells long, three to 
five cells broad at the base, the sinuses acute to lunulate, the 
margins entire; leaf cells thin-walled, the cell lumina angular- 
rounded, the trigones conspicuous, sometimes becoming coalesced, 
the cuticle faintly verruculose; the cells of the apical region 
mostly 24/i X 24/x, of the interior larger, those of the base to 
48/A X 24/A, a vitta not differentiated : underleaves distant, round- 
quadrate, entire, scarcely broader than the stem, attached in a 
straight line, 0.28 long and broad, the margins rounded, the cells 
20/i - 24/x long X 16/x broad, the cell walls uniformly thickened, 
especially along the margin: leaves of the flagelliform branches 
scale-like, ovate : sexual branches not seen. 

HABITAT : Not given. 

The distinguishing characteristics of the species are its golden 
brown color, and the almost filiform habit of the stems; the 
ascendent, more or less equally three-toothed leaves with large, 




FIG. 27. Bazzania tricuspidata (Steph.) Fulford. 1. Portion of a 
plant, ventral view, X 15. 2. Portion of a leaf and stem, dorsal view, X .30. 
3. A leaf, X 30. 4. A tooth of a leaf, X 310. 5. A cell from the apica 
portion of a leaf. X 400. 6. Underleaves, X 30. 7. Portion of the apical 
margin of an underleaf, X 310. Nos. 1-7 drawn from a portion of the type 
material. 



Bazzania 81 Grandistipulae 

thin-walled cells and conspicuous trigones; and the small, round- 
quadrate underleaves with entire margins and uniformly thickened 
cell walls. (FiG. 27, nos. 1-7). 

The plants are very small, and the leaves which are narrowed 
only a little from the base to the apex, are always ascendent and 
more or less equally three-toothed as seen in FIG. 27, nos. 1, 2 and 3. 
The teeth are sharply acute (FiG. 27, no. 4), and usually end in a 
two-celled point. The underleaves are always small, round-quadrate 
and entire. They are usually distant. The cells are mostly thick- 
walled, particularly along the margin (FiG. 27, no. 7). 

The species can readily be separated from B. diversicuspis, the 
only other small species of this Section, because of its golden brown 
color, the smaller size, the very acute teeth of the leaves, and the 
rounded underleaves with thick-walled cells, particularly along the 
margin. 

DISTRIBUTION : British Guiana: without locality, Quelch, the 
type (H). 

REFERENCES: Stephani (I cones, Mastigobryum no. 60). 

21. Bazzania longistipula (Lindenb.) Trevis. Mem. 1st. Lomb. 

13: 415. 1877. 

Mastigobryum longistipulum Lindenberg, in G. L. & N., Syn. Hep. 228. 1845. 
Mastigobryum c&nsanguineum Hampe & Lindenberg, in G. L. & N., op. cit. 

p. 717. 1847. 

Bazzania consanguine^, Trevis. op. cit. p. 414. 
Mastigobryum consanguineum var. brachyphyllum Stephani, Hedwigia 24: 

217. pi. 2, fig. 2. 1885. 

Mastigobryum brachyphyllum Gottsche in Stephani, loc. cit. (nomen nudum) 
Bazzania teretiuscula Spruce, Trans. & Proc. Bot. Soc. [Edinburgh] 15: 375. 

1885. Not M. teretiusculum Lindenberg & Gottsche. 
Bazzania longistipula var. Spruce ms., Hepat. Sprue. 
Mastigobryum paludosum Gottsche in Stephani, Hedwigia 25: 243. 1886. 

(nomen nudum) 
Mastigobryum phyllobolum Gottsche in Stephani, loc. cit. (nomen nudum) 

Not Bazzania phyllobola Spruce. 
Mastigobr^fum saxatile Gottsche in Stephani, Spec. Hep. 3: 470. 1908. (nomen 

nudum) 
Mastigobryum elegantulum Gottsche, Ann. Sci. Nat. Bot. Sen V. 1: 141. 1864. 

(nomen nudum); in Stephani, Hedwigia 25: 5. pi. 2. fig. 1-3. 1886. Not 

M. elegantulum I)e Notaris, 1874. 

Bazzania Krugiana Stephani, Hedwigia 27: 300. pi. 13. fig. 40. 1888. 
Mastigobryum Krugianum Stephani, Spec. Hep. 3: 447. 1908. 
Mastigobryum Hansenii Stephani, op. cit. p. 467. 
Bazzania longistipula var. polymastix Spruce ms., Hepat. Sprue. 
Bazzania decidua Spruce, Trans. & Proc. Bot. Soc. [Edinburgh] 15: 372. 1885. 
Mastigobryum deciduum Stephani, op. cit. p. 447. 
Mastigobryum Puiggarii Stephani, op. cit. p. 472. 

Plants in depressed mats or deep tufts, olive-green to golden 
brown or dark brown : stems slender to 6 cm. or more long, with 
leaves 1.5 mm. to 3 mm. broad, prostrate to ascending or erect; 
in longitudinal section the cells elongate, the medullary averaging 
0.16 mm. long, the cortical shorter, both averaging 18/A in diameter, 
the vertical walls uniformly thickened and containing frequent pits, 
the end walls thin: lateral branches 0.5 cm. to 1.5 cm. apart, diverg- 



M. Fulford 82 Bazzania 

ing at a wide angle : flagellif orm branches frequent, long : rhizoids 
colorless, present on some of the leaves of the flagelliform branches : 
leaf insertion curved in the upper part ; the leaves approximate to 
densely imbricated, ascendent, often deflexed and becoming 
wrapped around the stem when dry, unsymmetrically ovate, 0.8 
mm. - 2 mm. long, mostly 0.5 mm. - 1 mm. broad at the base, 
narrowed to the obliquely truncate, tridentate apex; the dorsal 
margin strongly convex from a rounded base which covers one-half 
to the entire axis and sometimes extends beyond, the ventral 
margin straight or slightly concave, the base often more or less 
expanded; the apex broad to narrow, tridentate, in slender forms 
sometimes bidentate or acute ; the teeth acute, two to six cells long 
and broad at the base, the acroscopic tooth the longest, the sinuses 
lunulate to deep and acute, the margins straight to undulate ; leaf 
cells thin-walled, the cell lumina angular-rounded, the trigones 
conspicuous, with convex sides, often becoming coalesced; the 
cuticle faintly verruculose; cells of the apical portion averaging 
24/i X 24/i, those of the median portion larger, those of the base 
36/A - 40/4 X 18/i, a vitta not differentiated : underleaves distant to 
imbricated, attached in a straight line, oblong-quadrate to quadrate, 
as broad or broader than the stem, 0.35 mm. - 1 mm. long X 0.35 
mm. - 0.7 mm. wide, often squarrose, the apex straight to rounded- 
entire, or faintly two to four undulate or lobed, the lateral margins 
straight to slightly convex, rounded at the bases, the cells similar 
to those of the apical portion of the leaf : leaves of the flagelliform 
branches scale-like, ovate, to 0.3 mm. long, acute to bifid : female 
branches occasional, one to several on a stem, the bracts and 
bracteoles similar; the outermost series ovate, to 0.35 mm. long, 
mostly bifid; the intermediate series larger, 0.8 mm. - 0.96 mm. 
long, 0.5 mm. wide, the margins crenulate, dentate to ciliate, the 
apical portion divided into two to four laciniae, four to six cells 
long, the cells uniform throughout, averaging 54/x X 18ft ; the inner- 
most series larger, one-third to one-fourth divided into usually 
three, dentate to ciliate laciniae, the lateral margins dentate to 
ciliate, the cells uniform, 45/x - 72/u, long X 18/i wide: perianth 
mouth laciniate, the laciniae crenulate, mostly three to five cells 
long: male branches and sporophyte not seen. 

HABITAT : In tufts or depressed mats on logs and soil in woods. 

The distinguishing characteristics of the species are its olive- 
green to yellow-brown color; the ascendent, obliquely truncated, 
tridentate leaves with thin cell walls, conspicuous trigones which 

FIG. 28. Bazzania langistipula (Lindenb.) Trevis. 1. Dorsal side of 
stem showing leaf attachment, X 30. 2. Portion of a stem, ventral view, 
X 12. 3. Leaves, X 30. 4. Cells from an apical tooth, X 260. 4a. A cell 
from the apical part of a leaf, X 350. 5. Cells from the dorsal margin of a 
leaf near the base, X 260. 6. Cells from the basal portion of a leaf, X 260. 
7. Underleaves, X 30. 8. Cells from the apical portion of an underleaf, 
X 260. 9. Portion of a transverse section of a stem, X 260. 10. Portion of 
a stem, ventral view, X 12. 11. Portion of a stem, ventral view, X 12. 
12. Portion of a stem, ventral view, X 12. 13. Underleaf, X 30. 14. Portion 
of a stem, ventral view, X 12. 15. Underleaf, X 30. Nos. 1-9 drawn from 
a portion of the type material, from St. Vincent; 10-11 from the type of 
B. teretiuscula Spruce, from Ecuador; 12-13 from the type of M. paludosum, 
from Guadeloupe; 14-15 from the type of M. saxatile, from Guadeloupe. 



M. Fulford 84 Bazzania 

often become coalesced, and angular-rounded cell lumina; and the 
round-quadrate to elongate underleaves which may be distant to 
densely imbricated. (FIGS. 28, nos. 1-15; 29, nos. 1-16). 

The plants show a high degree of variation in size and form 
of the leaves and underleaves. The variations in the underleaves 
are primarily in length and degree of convexity of the lateral 
margins, while those of the leaf are mostly in length, in the width 
of the apical portion, and in the size and shape of the teeth. 

STEPHANI (1886, 243) called attention to this variation among 
the plants. He stated that after an examination of Mastigobryum 
longistipulum from the HOOKER Herbarium (the type material), 
he considered it to be the normal form and M. consanguineum as 
the more lax mountain form. He added further, that Dr. 
GOTTSCHE'S Antilles species, M. paludosum with somewhat taper- 
ing leaves, M. brachyphyllum with distant, shorter underleaves, and 
M. phyllobolum G., a distorted form with shorter leaves, were 
variations of this species. He said that all these plants show the 
same leaf outline, strongly developed teeth, the uppermost of which 
is by far the longest, elongate underleaves, and the same angular, 
strongly thickened leaf tissue. They all grew erect in densely com- 
pact, deep tufts of a reddish brown color. He stated further that 
he was convinced that here was a whole series of forms of a wide- 
spread species, such as had been observed in other liverworts 
and mosses. To the above forms he later (1908, 470) added 
M. saxatile. 

An examination of the available material has led to the con- 
clusion that M. longistipulum, M. consanguineum, M. brachy- 
phyllum, B. teretiuscula Spruce, M. paludosum, M. phyllobolum G., 
M. saxatile, M. elegantulum, B. Krugiana, M. Hansenii, B. decidua, 
B. longistipula var. polymastix and M. Puiggarii are, in part, 
different expressions of the species under differing conditions of 
growth, a situation similar to that found in the more northern B. 
tricrentata (FULFORD, 1936). 

The plants of the type collection of M. longistipulum, from St. 
Vincent (see FIG. 28, nos. 1-9), are robust, but the stems are rarely 
over 6 cm. long and 3 mm. broad, and are copiously branched, with 
the branches diverging at a wide angle. The leaves are ascendent, 
the underleaves densely imbricated, and elongate, with rounded 
to faintly two- to four-lobed apices, and nearly straight sides from 
slightly rounded bases. 

Most of the plants of B. teretiuscula Spruce, from Ecuador, are 
very similar (FiG. 28, nos. 10-11), except that they are less robust 
and the branches occur less frequently, although a few of the stems 
are as well developed as those of M. longistipulum (compare nos. 
2 and 10). The underleaves are distant to imbricated, and while 
most of them are elongate with nearly straight sides, a few are 
shorter and have bulging sides. The apices often tend to be more 
deeply lobed. M. paludosum (FiG. 28, nos. 12-13) , from the Antilles 
is very similar but not so large. 



Bazzania 85 Grandistipulae 

Af. elegantulum (FiG. 29, nos. 6-12), collected in Jamaica ap- 
proximates B. teretirtscula in appearance, although the stems are 
not so long. The leaves are short, ascending, and the underleaves 
are mostly approximate and quadrate, with straight to bulging 
sides. A/, saxatile (FiG. 28, nos. 14-15), from Guadeloupe, M. 
Hansenii (FiG. 29, nos. 3-5), from Jamaica, and B. decidua (FiG. 
29, nos. 13-14), from Ecuador, are very similar. (B. decidua is 
lighter in color than most of the other plants, and the leaves and 
underleaves are well developed but distant. The stems are long and 
little branched.) B. Krugiana (FiG. 29, nos. 1-2), collected in 
Santo Domingo, differs only in that the underleaves are less well 
developed and are distant. A cross section of the stem shows that 
the walls of the stem cells are not so thick as in the other variations, 
but this too is a variable characteristic, dependent in part at least 
on the type of habitat in which the plant grows. 

A/. Puiggarii (FiG. 29, nos. 15-16), from Brazil, is rather small 
and yellow-brown, with the underleaves mostly four-lobed at 
the apex. 

STEPHANI'S figures (1885, 217-218. pi. 2, fig. 2.), of M. con- 
sanguineum var. brachyphyllum, from Guadeloupe, suggests that 
this plant is a variation similar to that shown by B. Kmgiana. His 
figures of M. consanguineum drawn from the type collected by 
MORITZ in Colombia show a plant with leaves little imbricated, and 
underleaves approximate, elongate, with nearly parallel margins, a 
plant with the characteristics of M. longistipulum but not so robust. 

The stems may be very slender to medium size and are usually 
branched. The poorly developed plants and the growing tips of 
the more robust forms are olive-green but most plants soon be- 
come strongly pigmented with brown. Those from the higher 
altitudes often take on a golden brown color. Some tend to grow 
in deep patches while others are found in depressed mats or 
scattered among mosses. 

The leaves may be distant to densely imbricated and are as- 
cendent, with the lower margins nearly straight. They may be 
nearly plane or deflexed and sometimes are tightly wrapped around 
the stem when dry. They are unsymmetrically ovate with a mostly 
obliquely truncate, tridentate apex. The length varies from 0.8 
mm. or less on poorly developed forms to as much as 1.5 mm. on 
robust stems. The teeth vary in shape and size but are usually 
strongly developed, with the acroscopic tooth the longest. The leaf 
cells are thin-walled, with conspicuous trigones. The size of the 
trigones varies with the age of the plant and its conditions of 
growth. They have bulging sides and when large may become 
coalesced. 

The underleaves also exhibit much variation. They may be 
distant to densely imbricated, and as wide or wider than the stem. 
The insertion forms a straight line. On many plants, most of 
them are rectangular in outline, with nearly straight lateral 
margins and an entire, faintly two- to four-lobed apex. On the 



M. Fulford 86 Bazzania 

same plant may also be found shorter underleaves, quadrate, with 
sometimes straight, but more often bulging sides. On some stems 
only this latter sort occur. Sometimes they are much reduced and 
very distant, and tend to be more deeply lobed or cut. 

The female branches are not common. The bracts and 
bracteoles are of the usual sort, ovate to oblong-ovate, with crenu- 
late to dentate margins, and divided at the apex into two to four 
laciniae. The perianths were not mature. The mouth is laciniate, 
with the laciniae three to five cells long (see FIG. 29, nos. 8-12). 

The species is distinguished from B. Breuteliana and B. acu- 
minata because of its much smaller size, its brown color, the as- 
cendent, obliquely truncate leaves which become conspicuously 
narrowed toward the apex, and its smaller underleaves. It differs 
from B. tricuspidata in its mostly larger size, the narrow, obliquely 
truncate leaf apices, and in the cells of the underleaf. Regardless 
of the shape of the underleaves of depauperate forms of B. longi- 
stipula the marginal rows of cells are never noticeably thick-walled, 
but instead are thin-walled with small but distinct trigones. 

DISTRIBUTION : Dominica: Morne Diablotin, Elliott 672, cited by 
Spruce (1895, 356). Guadeloupe: without locality, Hb. Gottsche, the 
type of M. paludosum (H); without locality, Hb. Gottsche, the type of M. 
mxatile (H); without locality, Parker (NY); Soufriere, Duss 136 (NY); 
Matouba, Duss 211 (NY); without locality, THerminier, Germain, cited by 
Stephani (1908, 470); without locality, THerminier, the type of M. consan- 
guinium var. brachyphyllum, cited by Stephani (1885, 217). Jamaica : 
without locality, Harris (NY) ; summit of Blue Mountain Peak, Maxon & 
Killip 1104 (Y, NY); Blue Mountain Peak, 6500-7325 ft., Underwood 1784 
(NY); Sir John's Peak, E. G. Britton 1197 (NY); without locality, Hansen, 
the type of M. Hansenii (H) ; without locality, Rheder, the type of M. 
elegantulum G. (H, B) ; New Haven Gap, Patterson (F). Puerto Rico: 
Rio de Maricao, E. G. Britton 2682 (Y). Santo Domingo (Dominican 
Republic): without locality, Eggers, the type of B. Krugiana (H, B). 
St. Vincent: without locality, Guilding (NY); without locality, Hooker 
Hb., the type of M. l&ngistipulum (H, V). T r i n i d ad : without locality, 
Beyrick, cited by G. L. & N. (1845, 231). - C ol om b i a : "Sierra Nevada, 
Prov. Merida", Moritz, the type of M. consanyuineum (H). Ecuador : 
Mt. Tunguaragua, Spruce, the type of B. teretiuscula Spruce, Hepat. Sprue. 
(Y, NY) ; the same locality, Spruce, the type of B. decidua, Hepat. Sprue. 
(NY). Peru: Mt. Campana, Spruce, the type of B. longistipula var. 
polymastix (NY). Bolivia: Yungas, Jay (Y). Brazil: Serra 

FIG. 29. Bazzania longistipula (Lindenb.) Trevis. 1. Portion of a 
stem, ventral view, X 30. 2. A cell from the apical portion of a leaf, X 400. 
3. Portion of a stem, dorsal view, X 30. 4. Portion of a plant, ventral view, 
X 30. 5. A cell from the apical portion of a leaf, X 400. 6. Portion of a 
plant, ventral view, X 30. 7. A cell from the apical portion of a leaf, X 400. 
8. Female bract of an intermediate series (immature), X 30. 9. A lacinia 
of a bract of this series, X 260. 10. Female bract of the innermost series 
(immature), X 30. 11. Portion of a lacinia of a bract of this series, X 260. 
12. Portion of the perianth mouth, X 260. 13. Portion of a plant, ventral 
view, X 15. 14. A cell from the apical portion of a leaf, X 400. 15. Portion 
of a plant, ventral view, X 15. 16. A cell from the apical portion of a leaf, 
X 400. Nos. 1-2 drawn from a portion of the type of B. Krugiana; 3-5 from 
a portion of the type of Af. Hansenii; 6-12 from a portion of the type of Af. 
elegantulum; 13-14 from a portion of the type of B. decidua; 15-16 from a 
portion of the type of M. Putggarii. 



M. Fulford 88 Bazzania 

do Mar, Dusen, the type of M. Puiggwrii (H); Apiahy, Puiggari, cited by 
Stephani (1908, 472). 

REFERENCES: Bescherelle (1893, 187); Hampe (1847, 328); Lindenberg 
& Gottsche (1851, 60; 88); Spruce (1895, 356); Stephani (1886, 243; 1901-05, 
98; 1903, 23; 1908, 447, 467, 470, 472; /cones, Mastigobryum nos. 56, 57, 58, 
163, 171 a, b, c, d, 173). 

22. Bazzania latidens (Gottsche) comb. nov. 

Maxtigobryum latidens Gottsche in Stephani, Hedwigia 25: 134. pi. 5. fig. 7-9. 
1886. 

Plants medium to large, golden yellow to yellow-brown, tinged 
with red in the older portions: stems to 5 cm. or more in length, 
with leaves to 4 mm. broad: lateral branches frequent, 1 cm. or 
more apart, diverging at a wide angle; flagelliform branches oc- 
casional ; rhizoid not seen : the line of leaf insertion curved in its 
upper part; the leaves subimbricated to imbricated, plane to de- 
flexed, unsymmetrically ovate to elongate, straight to ascendent, 
becoming a little falcate, mostly 2 mm. long, 0.8 mm. broad at the 
base, narrowed a little to the obliquely truncate, tridentate apex; 
the dorsal margin strongly arched from a cordate base, extending 
across the stem, the ventral margin concave, the base little dilated, 
the apex three-toothed, the teeth variable, mostly large, six to eight 
cells long, five to eight cells broad at the base, the sinuses deep, 
acute to lunulate, the margins entire ; the leaf cells thin-walled, the 
cell lumina angular-rounded, the trigones conspicuous, with convex 
sides, sometimes becoming coalesced, the cuticle verruculose; the 
cells of the apical region 20/i - 24/x in diameter, of the medium 
portion larger, and of the base 40/A - 48/x X 24/x, a vitta not differ- 
entiated: underleaves approximate to imbricated, rectangular in 
outline, attached in a straight line, broader than the stem, 0.6 
mm. - 0.9 mm. long, 0.55 mm. broad, the lateral margins nearly 
straight, entire, the apex truncate, entire, undulate to slightly two- 
to four-lobed, the cells tending to be in rows, mostly 16/x - 24/A 
X 16^i, the trigones conspicuous: sexual branches not seen. 

HABITAT : Over rocks. 

The distinguishing characteristics of the species are its medium 
to large size, the golden brown color, the plane to deflexed leaves 
which are elongate, spreading, and strongly three-toothed, with 
cells 20/x - 24/t in diameter, thin-walled and with conspicuous 
trigones; and the rectangular underleaves with nearly straight, 
entire lateral margins, and truncate, entire, undulate to faintly 
lobed apices. (FiG. 30, nos. 1-6). 

The type in the STEPHANI Collection at Harvard consists of one 
robust stem about 4 cm. long. The outstanding characteristics of 
the plant are its elongate leaves, and rectangular underleaves which 
are only a little curved at the base, and are entire to faintly two- to 
four-lobed at the apices. 

Plants with similar characteristics were collected by UNDER- 
WOOD on Blue Mountain Peak in Jamaica. These plants grew in 
mats or tufts and are light green at the growing tips and golden 




FIG. 30. Bazzania latidens (Gottsche) Fulford. 1. Portion of a 
plant, ventral view, X 15. 2. A leaf, X 15. 3. A tooth of a leaf, X 310. 
4. A cell from the apical portion! of a leaf, X 400. 5. An underleaf, X 30. 
6. A portion of the apical margin of an underleaf, X 310. Nos. 1-6 drawn 
from a portion of the type material. 



M. Fulford 90 Bazzania 

brown tinged with red in the older portions. The leaves are not 
quite so long but are of the same shape, with similar teeth and cell 
configuration. The underleaves are long-rectangular, with the cells 
arranged more or less in rows as in the type. On some of the 
stunted plants in which the internodes are very short, the under- 
leaves tend to be a little shorter, but otherwise the characteristics 
are the same. 

The species has spreading leaves and elongate underleaves with 
apices truncate and three- or four-lobed, similar to those of B. 
Breuteliana. However, the yellow-green to brown color, the well 
defined teeth of the leaves, and the larger trigones of the leaf -cells 
of B. latidens are usually sufficiently distinctive to easily 
separate the two. 

The species is perhaps more closely related to B. longistipula 
and is intermediate between that species and B. longa. B. latidens 
is always larger than J5. longistipula and the leaves are usually 
elongate and spreading. The leaves of B. longistipula are always 
ascendent, and very obliquely truncate at the apices. The under- 
leaves of B. latidens are always elongate, with the lateral margins 
more or less parallel. This characteristic is also present in the 
robust forms of B. longistipula. 

DISTRIBUTION: Jamaica: Blue Mountain Peak, 6500-7325 ft, Under- 
wood 1483, 1484, 1490, 1512 (NY); Cinchona, Harris 11, 135a (NY). 
Brazil : without locality, Glaziou 1792, the type (H). 

REFERENCES: Stephani (1909, 529; /cones, Mastigobryum no. 390). 

23. Bazzania longa (Nees) Trevis. Mem. 1st. Lomb. 13: 415. 1877. 

Jungermannia longa Nees, Linnaea 6: 623. 1831. 

Jungermannia stolonifera Sieb. PI. crypt, exsicc. n. 35. Linnaea, loc. cit. Not 

J. stolonifera Swartz. 

Mastigobryum longum Nees, in G. L. & N., Syn. Hep. 231. 1845. 
Mastigobryum Gottscheanum Lindenberg, op. cit. p. 224. 
Bazzania Gottscheana Trevisan, op. cit p. 414. 
Mastigobryum speciosum Gottsche, Husnot PI. Antilles no. 213, 1868. Husnot, 

Rev. Bryol. 2: 3. 1875. Gottsche in Stephani, Hedwigia 25: 233. pi. 1. fig. 

1-3. 1886. 

Mastigobryum subfalcatum Gottsche in Stephani, op. cit p. 234, pi. 1. fig. 4-6. 
Bazzania subfalcata, Spruce, Jour. Linnean Soc. Bot. 30: 356. 1895. 
Mastigobryum tenue Stephani, Spec. Hep. 3: 448. 1908. 
Mastigobryum trinitatis Stephani, Spec. Hep. 6: 483. 1924. 

Plants medium to robust, dark green becoming deep red-brown : 
stems stout, to 10 cm. long, with leaves to 5.5 mm. broad, ascend- 
ing to erect; in longitudinal section the cells elongate, averaging 
0.16 mm. long, the cortical shorter, both averaging 16/x in diameter, 
the vertical walls strongly thickened and containing frequent pits, 
the end walls thin, the cells of the outer layers strongly pigmented : 
lateral branches distant, diverging at a wide angle: flagelliform 
branches frequent, long: rhizoids numerous, present on the bases 
of the leaves of the flagelliform branches : the line of leaf insertion 
curved in its upper half; leaves imbricated, strongly falcate, de- 




FIG. 31. Bazzania longa (Nees) Trevis. 1. Portion of leaf and stem, 
dorsal view, X 30. 2. Portion of plant, ventral view, X 12. 3. Leaf, X 30. 
4. Cells from an apical tooth of a leaf, X 260. 5. A cell from the apical 
portion of a leaf, X 350. 6. Underleaves, X 30. 7. Female bract of the outer- 
most series, X 30. 8. Female bracts of intermediate series, X 30. 9. One 
of the laciniae of a female bract of the innermost series (immature), X 100. 
Nos. 1-9 drawn from the type material. 



M. Fulford 92 Bazzania 

flexed, often becoming connivent when dry, unsymmetrically ovate, 
often elongate, 1.5 mm. - 2.5 mm. long, to 1.5 mm. broad at the 
base, narrowing to the obliquely truncate, tridentate apex ; the dor- 
sal margin strongly convex from a rounded base which extends 
across the stem, the ventral margin concave, the base a little dilated ; 
the teeth large, unequal, the acroscopic tooth longer than the others, 
four to eight cells long, three to five cells broad at the base, the 
sinuses deep, acute to rounded, the margins straight to repand; 
leaf cells thin-walled, the cell cavities stellate, the trigones very 
large, with strongly convex sides, soon becoming coalesced by the 
deposition of secondary thickenings, the cuticle verruculose; cells 
of the apical portion and dorsal lobe 24/x - 34/x X 24/x, of the median 
portion larger, and of the base 54^ X 18/x, a vitta not differentiated : 
underleaves distant to imbricated, the line of attachment straight, 
round-quadrate to elongate, 0.45 mm. - 1.2 mm. long, 0.45 mm. - 0.8 
mm. broad, the lateral margins entire, often convex from a rounded 
base, the apex rounded, entire, to faintly two- or four-lobed: 
leaves of the flagelliform branches small, ovate, to 0.28 mm. long, 
spreading, the apex acute to shortly bifid, the cells very large: 
female branches occasional, solitary, one to several on a stem, the 
bracts and bracteoles similar, ovate; the outermost series short, 
rounded to shortly bifid; the intermediate series longer, shortly 
trifid, the teeth and margins mostly entire ; the innermost series to 
1.25 mm. long, to one-seventh divided into three short, crenate 
laciniae, the cells to 48/x long, 18/x wide, thick-walled, the lateral 
margins crenate to short spinose: perianths to 6 mm. long, the 
mouth short-spinose, the cells thick-walled : male branches not seen. 

HABITAT : On rocks, logs and over tree bases. 

The distinguishing characteristics of the species are its large 
size and dark red-brown color; the large, mostly falcate, obliquely 
truncate, unequally tridentate leaves with large teeth, large cells, 
stellate cell-cavities, and very large trigones ; and the subquadrate 
to elongate underleaves which have convex to straight lateral 
margins and rounded or faintly lobed apices. (FlGS. 31, nos. 1-9; 
32, nos. 1-16; 33, nos. 1-12). 

The plants of /. longa (FiG. 31, nos. 1-9), collected by SlEBER 
in Martinique, are robust, to 10 cm. or more in length, and 4 mm. 
or more broad, and are unbranched. They are light to dark green 
at the growing tips but are characteristically a deep red-brown in 
the older portions. The leaves are long, unsymmetrically ovate, 
slightly to strongly falcate, and narrowed to the tridentate apices. 
The teeth are mostly large, often spreading, with the acroscopic 
tooth often the longest. The cell walls are at first thin, but soon 
become thickened by the deposition of secondary thickenings, and 
the trigones are large and very often coalesced. The cell lumina 
are stellate as shown in FIG. 31, nos. 4 and 5. The underleaves are 
distant, quadrate to elongate, broader than the stem, with the 
lateral margins entire and more or less parallel from a curved 
base, and the apices entire or faintly two- to four-lobed. 

The plants of the type collection of M. Gottscheanum (FiG. 32, 
nos. 13-16) are slightly smaller, to 6 cm. long, dark red-brown, 



oc 

OS 
o o /C 




FIG. 32. Bazzania longa (Nees) Trevis. 1. Portion of a stem, ventral 
view, X 12. 2. Portion of a stem, dorsal view, X 30. 3. Leaf, X 30. 4. Leaf 
apices, X 30. 5. Apical tooth, X 260. 6. A cell from the apical portion of a 
leaf, X 350. 7. Cells from the dorsal margin of a leaf, X 260. 8. Cells from 
the basal portion of a leaf, X 260. 9. Underleaves, X 30. 10. Cells from the 
upper margin of an underleaf, X 260. 11. Portion of a transverse section of 
a stem, X 260. 12. Male bract, X 30. 13. Portion of a stem, ventral view, 
X 12. 14. Portion of a stem, dorsal view, X 30. 15. A cell from the apical 
portion of a leaf, X 350. 16. Underleaves, X 30. Nos. 1-12 from a portion 
of the type of M. tenue; 13-16 from a portion of the type of M. Gottscheanum. 



M. Fulford 94 Bazzania 

and the leaves are deflexed and usually strongly falcate, although 
some leaves are only slightly so. The leaf cells are large, with very 
large, rounded trigones and stellate cell lumina. The underleaves 
are round-quadrate, with the lateral margins entire, and the apices 
rounded or faintly lobed. The plants of the type of M. tenue (FlG. 

32, nos. 1-12) are also red-brown, approximately the same size, but 
most of the leaves are not quite so falcate, and many of the under- 
leaves are longer. The plants of M. subfcUcatum (FlG. 33, nos. 1-6) 
are also large and reddish brown, with deflexed, falcate leaves and 
mostly round-quadrate underleaves. They are identical with those 
of J. longa. 

Most of the plants of the type collection of M. speciosum (FlG. 

33, nos. 7-8) are very large, to 10 cm. long and 4 mm. broad, a 
deep red-brown to black color, with the underleaves elongate rather 
than round-quadrate. In the other characteristics, leaf insertion, 
leaf shape, teeth, trigones, and the margins of the underleaves, the 
plants agree with those described above. Smaller plants in the 
same collection are intermediate or practically identical with M. 
Gottscheanum, while the larger ones are typical of large plants of 
J. longa. The female bracts of plants collected by ELLIOTT in 
Dominica, no. 1080, are identical with those of the type of M. 
subfalcatum. 

M. trinitatis (FlG. 33, nos. 9-10) collected by CRtteER in Trini- 
dad must also be included here. The plants are rather small, the 
pigmentation is not so strongly developed, and many of the leaves 
are not so falcate, but the coloration, type of leaf, and cell pattern 
are the same. It is obvious that the plants are poorly developed. 

In some instances the above species have been identical, in 
others they have differed in size, leaf curvature, degree of pig- 
mentation, length of underleaves, etc., with much variation in the 
individuals of one mat, so that no definite limits of variation can 
be assigned to any one of them. They appear to represent mostly 
habitat variations of one unit, J. longa. 

var. papillata (Steph.) comb. nov. 

Mastigobryum papillatum Stephani, Spec. Hep. 3: 526. 1909. 

This variety, collected in Dominica (FlG. 33, nos. 11-12), differs 
from the species in that the cuticle of the leaves and underleaves 
is very conspicuously roughened by large, coarse, wart-like pro- 
tuberances. The cuticle of J. longa and of the synonyms described 
above varies from faintly to strongly verruculose, but in no 
instance are the verruculae even half so large as those of the variety. 

DISTRIBUTION: Dominica : without locality or collector's name (NY); 
Morne Diablotin, Elliott 678, cited by Spruce (1895, 356) ; without locality, 
Elliott 1080 (G); without locality, Elliott, the type of M. papillatum (H) ; 
without locality, Elliott, as M. arcuatum (G). Cuba : without locality, 
Wright, Hep. Cubensis Wrightianae, as M. brasiliense (H). Guade- 
loupe: without locality, I'Herminier (NY); without locality, 1'Herminier, 
the type of AT tenue (H) ; without locality, Richard (NY); without locality, 




FIG. 33. Bazzania longa (Nees) Trevis, 1. Portion of a stem, ventral 
view, X 15. 2. A cell from the apical portion of a leaf, X 400. 3. A female 
bract, outermost series, X 30. 4. A female bract, innermost series, X 30. 
5. A portion of an apical tooth of a bract of this series, X 300, 6. A portion 
of the perianth mouth, X 310. 7. Portion of a plant, ventral view, X 16. 
8. A cell from the apical portion of a leaf, X 400. 9, Portion of a plant, 
ventral view, X 15. 10. A cell from the apical portion of a leaf, X 400. 
B. longa var. papillata (Steph.) Fidford. 11. Portion of a plant, ventral 
view, X 15. 12. A cell from the apical portion of a leaf, X 400. Nos. 1-6 
drawn from a portion of the type of Af. subfalcatumi 7-8 from a portion of 
the type of M. speciosum; 9-10 from a portion of the type of 3/. trinitatis; 
11-12 from a portion of the type of A/. papUlatum, 



M. Fulford 96 Bazzania 

FHerminier, the type of M. subfalcatum (G, H) ; Morne-PEchelle, Duss 394 
(NY); Soufriere, Duss 83 (NY). Jamaica : John Crow Peak, Under- 
wood 680 (NY) ; Tweedside, Underwood 2042 (Y). Martinique: with- 
out locality, Sieber 35, the type (H, NY) ; Mt. Pelee, Husnot, PL Antilles 213, 
the type of M. speciosum (G) ; Mt. Pelee, Duss 394, 395, 584, 606 (NY); 
without locality, Bordaz, cited by Stephani (1908, 469) ; without locality or 
collector's name (H). Puerto Rico: Sierra de Naguabo, Shafer 3750 
(NY); the same, Johnson 1620 (NY). St. Kitts: without locality, 
Breutel, the type of Af. Gottscheanum (B) ; without locality, Breutel (NY). 
Trinidad: without locality, Criiger, the type of M. trinitatis (G) ; 
without locality, Criiger (NY); without locality or collector's name, Hb. 
Jack (G). 

REFERENCES: Montagne (1843, 253); Lindenberg & Gottsche (1851, 66, 
105) ; Stephani (1903, 22; 1908, 468, 472; 1909, 519, 520; /cones, Mastigobryum 
nos. 59, 162, 170, 179, 396, 399, 400); Spruce (1895, 356). 

24. Bazzania jamaicensis (Lehm. & Lindenb.) Trevis. Mem. 1st. 

Lomb. 13: 414. 1877. 

Herpetium jamaicense Lehmann et Lindenberg, in Lehmann, Pug. PI. 7: 7. 

1838. 
Mastigobryum jamaicense Lehm. et Lindenb., in G. L. & N., Syn. Hep. 223. 

1845. 

Mastigobryum chamaecardion Herzog rns. (nomen nudum) 
Bazzania jamaicensis var. chamaecardia Herzog, Rev. Bryol. et Lichen. 11 : 

19. 1938. 
Mastigobryum aaracanum Stephani, Spec. Hep. 3: 525. 1909. 

Plants large, in deep tufts or mats, olive-green to brownish 
green, strongly pigmented with brown: stems stout, to 10 cm. or 
more long, with leaves 4 mm. broad, prostrate to suberect; stem 
cells in longitudinal section elongate, averaging 0.2 mm. long, the 
cortical shorter, both averaging 18/i in diameter, the vertical walls 
strongly thickened and containing frequent pits, the end walls thin ; 
lateral branches frequent, mostly more than 5 mm. apart, diverg- 
ing at a wide angle; flagelliform branches frequent, long, often 
branched; rhizoids colorless, present on the bases of some of the 
leaves of the flagelliform branches: the leaf insertion curved in 
its upper part; leaves imbricated, convex, strongly deflexed even 
when moist, unsymmetrically ovate, a little falcate, to 2 mm. long, 
1 mm. - 1.5 mm. broad at the base, narrowing to the rather broad, 
transversely truncate, tridentate, apex; the dorsal margin arched 
from a strongly rounded base, extending the width of the stem, the 
ventral margin concave, the base a little dilated, the apex irregu- 
larly three-toothed, the teeth acute, narrow to broad, two to six 
cells long, two to nine cells broad, some obscure, the sinuses narrow, 
acute to lunulate, the margins entire, straight to undulate ; the leaf 
cells thin-walled, the cell lumina angular-rounded, the trigones 

FIG. 34. Bazzania jamaicensis (Lehm. & Lindenb.) Trevis. 1. Portion 
of leaf and stem, dorsal view, X 30. 2. Portion of plant, ventral view, X 10. 
3. Leaf, X 30. 4. A cell from the apical portion of a leaf, X 350. 5. Cells 
of an apical tooth, X 260. 6. Cells from the dorsal base of a leaf, X 260. 
7. Underleaves, X 30. 8. Cells from the apical portion of an underleaf, X 260. 
9. Portion of a cross-section of the stem, X 260. 10. Male bracteoles, X 30. 
11. Male bracts, X 30. 12. Cells from the apical portion of a male bract, 
X 260. Nos. 1-9 drawn from a portion of the type material; 10-12 drawn 
from plants collected in Peru by LECHLER. 



M. Fulford 98 Bazzania 

small, conspicuous, with convex sides, soon becoming coalesced in 
the marginal and apical regions, the cuticle smooth to faintly 
verruculose; cells of the apical region and dorsal base averaging 
20/i X 20/i, those of the median portion a little larger, those of the 
base 45/i - 54/i X 27ft: underleaves approximate to imbricated, 
broader than the stem, attached in a straight line, appressed, 
broadly ovate from an auricled base, very concave when seen from 
above, 0.65 mm. long X 0.8 mm. - 1 mm. broad at the base, the 
margin entire; the cells of the interior 22/i - 32/x long and broad, 
the cell lumina angular-rounded, the trigones small, conspicuous, 
the walls thin, soon becoming thick by deposits of secondary thick- 
enings and coalesced trigones, the marginal rows slightly larger, 
the walls more uniformly thickened, deeply pigmented with brown : 
leaves of the flagelliform branches ovate, to 0.32 mm. long, mostly 
entire: male branches one to several on a stem; the bracteoles 
small, oblong, averaging 0.56 mm. long X 0.28 mm. wide, some- 
what concave from above, the apex rounded or with one or two 
obscure teeth ; the bracts larger, ovate, to 0.56 mm. long - 0.64 mm. 
broad at the base, the apex entire, or with two or three short teeth ; 
antheridia occurring singly: female branches and sporophytes 
not seen. 

HABITAT : On tree trunks. 

The species is distinguished by its large size, the olive-green 
color and brown pigmentation; the unsymmetrically ovate, trans- 
versely truncate, irregularly tridentate leaves; and the inflated, 
entire, ovate, auricled underleaves with dark borders which are 
appressed to the stem. (FiGS. 34, nos. 1-12; 35, nos. 1-4). 

B. jamaicensis is one of the easiest of the American species to 
recognize. The plants are robust, dark green with some brown 
pigmentation, and usually grow in prostrate mats. The leaves are 
large, broadly ovate, and the teeth while conspicuous, are short 
and broad. The underleaves offer the best diagnostic character- 
istics. They are broadly ovate from a cordate base. They are 
strongly concave when seen from above, and the margin, which is 
entire, is tightly appressed to the stem, so that they appear to be 
inflated. The marginal cells are often a little larger than the others, 
and become pigmented with brown or red-brown, the degree of 
coloring apparently dependent on the habitat. The underleaves of 
robust plants always have the borders deeply pigmented. 

M. caracanum, from Brazil, is not quite so robust as H. 
jamaicense from Jamaica, but in other respects is identical with it. 

B. jamaicensis var. chamaecardia, from Costa Rica, is much 
less robust. The stems are long, and the leaves and underleaves 
are distant. The concavity of the underleaves is not nearly so 
conspicuous, nor is the pigmentation of the marginal rows of cells 
so well developed. Since however, many intermediate stages be- 
tween this form and the more robust type occur, it would seem that 
the variety is probably the result of the special conditions under 
which it grew. 

The underleaves will serve to distinguish the species from all 
of those already described. It might possibly be confused with B. 
Wrightii, a description of which follows. 



wv 

d~\*J 




FIG, 35, Bdzzania jamicemis (Lehm, 4 Lindenb.) Trevis, 1, Por- 
tion of a plant, ventral view, X 15. 2, A cell from the apical portion of a 
leaf, X 400. 3. Portion of plant, ventral view, X 15. 4. A cell from the 
apical portion of a leaf, X 400, Nos. 1-2 drawn from a portion of the type 
of M. cflracanum from Brazil; 3-4 from a portion of the type of M, chamae- 
cardion from Costa Rica, 



M. Fulford _100 Bazzania 

DISTRIBUTION : Jamaica: without locality, Swartz, cited by Stephani 
(1909, 525); without locality, Hart, cited by Boswell (1875, 50); without 
locality or collector's name, Hb. Hooker, the type (NY) ; Blue Mountain Peak, 
7300 ft., Harris 20, 11060 (Y, NY) ; summit of Blue Mountain Peak, Under- 
wood 2505 (NY); slopes of Sir John, E. G. Britton 1209 (NY); Blue Moun- 
tain, Evans 239 (Y). Costa Rica: Cerro de las Lajas, Standley & 
Valerio 49949, 50713, 51446, 51643a, 51645, 52117, 52267, the type of var. 
chamaecardia (W) ; Segovia, Oersted, cited by Hampe (1851, 302).- 
Brazil : Rio de Janeiro, Guadichaud (NY); Caracas, Vainio, the type of 
M. caracanum (H). Ecuador: Cayambe, Jameson (NY). Peru : 
Tatanara, Lechler 3112 (NY). Venezuela: without locality, Fendler, 
cited by Stephani (1909, 525); Caracas, Funck & Schlim (NY). 

REFERENCES: Montagne (1844-46, 244); Hampe (1851, 302); Lindenberg 
& Gottsche (1851, 52); Boswell (1887, 50); Stephani (1909, 525; /cones, 
Mastigobryum nos. 380, 388). 

25. Bazzania Wrightii (Gottsche) comb. nov. 

Mastigobryum Wrightii Gottsche in Stephani, Hedwigia 25: 237. pi. 2, fig. 

24-26. 1886. 
Mastigobryum stoloniferum var. cubense Gottsche ms., Hep. Cub. Wrightianae. 

Plants medium size to large, light yellow-green, becoming 
deeply pigmented with yellow-brown in the older portions: stems 
slender to robust, 5 cm. or more in length, with leaves to 4.5 mm. 
broad, prostrate to ascending: stem cells in longitudinal section 
elongate, to 0.17 mm. long, the cortical shorter, both averaging 18/i 
in diameter, the vertical walls uniformly thickened and containing 
frequent pits, the end walls thin ; lateral branches frequent, diverg- 
ing at a wide angle ; flagellif orm branches numerous, long : rhizoids 
colorless, from the basal cells of the leaves of the flagelliform 
branches : the line of leaf insertion curved in its upper part ; leaves 
distant to imbricated, falcate, often becoming deflexed when dry, 
unsymmetrically ovate, 1.5 mm. - 3 mm. long, 1 mm. - 1.5 mm. 
broad at the base, narrowed a little to the more or less obliquely 
truncate, tridentate apex ; the dorsal margin arched from a curved 
base, covering one-half or more of the stem, often extending beyond, 
the ventral margin concave, the base dilated, the apex irregularly 
three-toothed, the teeth mostly small, acute, to six cells long, three 
to five cells broad at the base, the sinuses shallow, lunulate, the 
margins entire; leaf cells thin- walled, the cell lumina angular- 
rounded, the trigones conspicuous, with convex sides, soon becoming 
coalesced through the deposition of secondary thickenings, the 
cuticle faintly verruculose; cells of the apical portion and dorsal 
base mostly 24/t X 24/x, those of the median portion larger, those 
of the basal area 32/z - 48/i X 24/x, a vitta not distinct : underleaves 
distant to imbricated, subquadrate to orbicular, broader than the 
stem, attached in a straight line,. 0.5 mm. - 1.2 mm. long, 0.5 mm. - 
0.95 mm. broad at the base, the lateral margins convex from a 
rounded to cordate base, entire, the apical portion rounded, entire, 
straight to four-lobed, a little squarrose, the cells 24/4, - 32u, thin- 
walled, the trigones large, often coalesced, intermediate thickenings 
developed in the elongate cells of the apical portion: leaves of the 
flagelliform branches scale-like, ovate, acute to shortly bifid, spread- 
ing ; female branches occasional, solitary, one to several on a stem, 
the bracts and bracteoles similar, the outermost series small, bluntly 




FIG. 36. Bazzania Wrightii (Gottsche) Fulford. 1. Portion of a 
plant, ventral view, X 15. 2. Portion of a stem and a leaf, dorsal view, X 30. 
3. A cell from the apical portion of a leaf, X 400. 4. A tooth of a leaf, X 310. 
5. Cells from the apical margin of an underleaf, X 310. 6. Portion of a plant, 
ventral view, X 15. 7. Portion of a leaf and stem, dorsal view, X 15. 8. A 
cell from the apical portion of a leaf, X 400. 9. Ventral bases of leaves 
attached to the stem, X 15. 10. Cells from the apical margin of an under- 
leaf, X 310. 11. Female bract of an outer series, X 30. 12. Female bract of 
an intermediate series, X 30. 13. One of the laciniae of a bract of the inner- 
most series, X 310. 14. Portion of the mouth of the perianth, X 100. Nos. 
1-5 drawn from a portion of the type material; 6-14 from a portion of the 
type of M. stoloniferum var. cubense Gottsche, from Cuba. 



M. Fulford 102 Bazzania 

ovate to shortly bifid; the intermediate series longer, shortly bi- 
trifid, the lateral margins entire; the innermost series larger, to 
one-sixth divided into two or three crenulate laciniae, the lateral 
margins crenulate to short ciliate, the cells to 48/* long, 24/t wide, 
with irregularly thickened walls: perianth (immature) mouth long 
laciniate-ciliate : male branches not seen. 

HABITAT : On logs and tree bases. 

The distinguishing characteristics of the species are its size 
and yellow- to brownish green color ; the more or less falcate leaves 
with short teeth and shallow sinuses; the thin- walled leaf cells, 
with conspicuous trigones which soon become coalesced, and 
angular-rounded cell lumina; and the subquadrate to orbicular 
underleaves, attached in a straight line, with lateral margins con- 
vex from a rounded to cordate base, and apices rounded to undulate 
or lobed and made up of rather large, elongate cells with thin walls, 
large trigones, and intermediate thickenings. (FiG. 36, nos. 1-14). 

The plants of the type collection (FiG. 36, nos. 1-5) are greenish 
yellow-brown in color, the stems are large, and with leaves 3 mm. 
or more broad, but they are for the most part, quite fragile and 
break readily. The leaves are approximate to imbricated and the 
underleaves are distant or approximate. The underleaves are 
rounded at the bases, and the mostly rounded apical margins very 
often show a deeper red-brown pigmentation than the interior 
or base. 

The only difference between these plants and those of M. 
stoloniferum var. cubense (FiG. 36, nos. 6-14), also collected in 
Cuba by WRIGHT, is that the latter are more robust, mostly but not 
always, light yellow-green in color, they are not brittle, and the 
underleaves are larger and densely imbricated. The bases of the 
underleaves are mostly strongly cordate instead of rounded as in 
M. Wrightii, but I believe that this is a characteristic of the species 
in its luxuriant form. The leaf shape, the teeth, cell size, walls 
and trigones, as well as the cell configuration, including the pres- 
ence of intermediate thickenings in the larger, elongate cells of the 
underleaves are identical and the two should therefore, I believe, 
be considered as different expressions, perhaps due to habitat 
conditions, of one species. 

The species is somewhat similar in several respects to B. 
jamaicensis. The two species differ in that the leaves of B. Wrightii 
are falcate, and the underleaves while often somewhat concave 
when seen from above, never appear to be inflated, and the 
margins are never closely appressed to the stem. The presence of 
intermediate thickenings in the elongate cells of the apical portion 
will also aid in separating the two. 

DISTRIBUTION: Cuba : Mt. Verde, Wright, Hep. Cub., the type (H, Y, 
NY) ; without locality, Wright, Hep. Cub., as M. stoloniferum var. cubense 
(H, Y, NY); Loma del Gato, Fr. Cl&nent 351 (NY). J amaica : Morse's 
Gap, Evans 48 (Y) ; Tweedside, Underwood 2042 (NY). Puerto Rico: 
without locality, Sintenis 3, 18, 33, 62, 84, 124, 130, cited by Stephani 
(1888a, 279). 

REFERENCES: Stephani (1888a, 279; 1909, 524; Icones, Mastigobryum 
no. 408). 



Section 2. Connatae 

This Section includes plants with more or less three-toothed 
leaves which are entire to more or less serrulate or spinose along 
the margins in the apical part. The underleaves are entire to 
variously and irregularly incised, serrate and toothed along the 
margins, and may have a hyaline border of several rows of cells 
or may be chlorophyllose throughout. The underleaves are at- 
tached in a straight line and are connate with the ventral base of 
one or a pair of leaves. This attachment with the leaf is often not 
conspicuous except under high magnification. 

The American species of this Section show three distinct tend- 
encies of development within the group. One of these, exemplified 
by only one species, B. Fendleri from Brazil, Ecuador and 
Venezuela, has leaves only faintly three-lobed, with the margins 
entire, and the cells thick- walled and with large trigones; the 
underleaves are round-quadrate, faintly four-lobed, with entire 
margins, and connate with one leaf. 

A second tendency is to be found in the species of the West 
Indies. The five species are very closely related and show a con- 
siderable degree of variation in their characters. The leaves vary 
from obscurely to strongly three-toothed ; from entire to serrulate 
or spinose in the apical region ; and the underleaves may be entire 
to sharply dentate or spinose, with a hyaline border, or chloro- 
phyllose throughout. All are connate by a few cells with one leaf. 
The distribution of some of the species is limited to one or two 
of the islands. 

The third tendency appears in the group of species which occurs 
south of the Equator from Peru to the Straits of Magellan with 
some of the species or their near relatives also in South Africa, 
Australia and New Zealand. This group is characterized by hav- 
ing the underleaves connate with a pair of leaves. The leaf apices 
may be obscurely to strongly three-toothed, entire to spinose, and 
the underleaves which are usually spinose-dentate have hyaline 
borders or are chlorophyllose throughout. 

Key to the Species 

1. Underleaves connate with a pair of leaves. 

2. Hyaline cells of the underleaves forming a conspicuous, broad border, 
on at least some of the underleaves . . .32. B. peruviana (p. 119) 
2. Hyaline cells of the underleaves restricted to a marginal row, or scat- 
tered, or absent; the cuticle noticeably verruculose, 

33. B. Skottsbergii (p. 122) 

1. Underleaves inconspicuously connate by a few (often only one or two) 
cells with one leaf only. 
2. Underleaves chlorophyilose throughout. 




FIG. 37. Bazzania Fendleri (Steph.) Fulford. 1. Portion of a plant, 
ventral view, X 15. 2. Portion of a leaf and stem, dorsal view, X 30. 3. A 
leaf, X 15. 4. A cell from the apical portion of a leaf, X 400. 5. Cells from 
the basal portion of a leaf, X 310. 6. Underleaf, X 15. Drawn from the 
type material. 



Bazzania 105 Connatac 

3. Underleaves entire or nearly so. ... 26. B. Fendleri (p. 105) 
3. Underleaves variously incised and toothed along the margin, 

31. R cubensis (p. 116) 
2. Underleaves bordered by a band of hyaline cells. 

3. Leaves conspicuously three-toothed, the margins strongly serrate 

and spinose 30. B. armatistipula (p. 114) 

3. Leaves conspicuously three- toothed, the margins obscurely ser- 
rate 29. B. Eggersiana (p. 112) 

3. Leaves inconspicuously three-toothed to truncate-undulate, the 
margins strongly serrate and spinose. 28. B. pycnophylla (p. 110) 
3. Leaves mostly rounded-entire, the apical margins only serrate, 

27. B. Schwaneckiana (p. 106) 

26. Bazzania Fendleri (Steph.) comb. nov. 
Mastigobryum Fendleri Stephani, Spec. Hep. 3: 436, 468. 1908. 

Plants medium to large, light greenish brown, more strongly 
pigmented in the older portions : stems to 5 cm. long, with leaves 
to 3.5 mm. broad, prostrate to ascending; lateral branches 5 mm. 
or more apart, diverging at a wide angle; flagelliform branches 
numerous, long ; rhizoids on the bases of the leaves of flagelliform 
branches : the line of leaf insertion little curved in its upper part ; 
the leaves imbricated, convex, ascendent, a little deflexed when dry, 
unsymmetrically ovate, to 1.5 mm. long, 0.7 mm. broad at the base, 
narrowed a little to the more or less transversely truncate, obscurely 
tridentate apex; the dorsal margin convex from a curved base, 
extending half way across the stem, the ventral margin nearly 
straight, the base scarcely dilated, the apex irregularly and ob- 
scurely one-, two-, or three-toothed, the teeth one to three cells 
high, two or three cells broad at the base, the sinuses lunulate, the 
margins entire; leaf cells thick- walled, the trigones large, with 
irregular, rounded sides, often becoming confluent, intermediate 
thickenings sometimes present, the cell lumina angular-rounded, 
the cuticle verruculose; cells of the apical portion averaging 16/* 
in diameter, those of the interior larger, those of the base 32/u, 
X 16/i, a vitta not differentiated: Underleaves large, imbricated, 
broader than the stem, round-quadrate, connate with one leaf, the 
line of attachment on the stem a straight line, 0.9 mm. broad and 
long, narrowed a little to the base, the margins recurved, the 
lateral margins convex, entire, the apical margin entire to faintly 
four-lobed, the cells as in the leaf: leaves of the flagelliform 
branches scale-like, to 0.2 mm. long, acute to shortly bifid: sexual 
branches not seen. 

HABITAT : Not given. 

The distinguishing characteristics of the species are its size, 
and light greenish brown color ; the obscurely two- or three-toothed 
leaves with thick-walled cells with angular-rounded lumina, and 
large trigones with irregular, convex sides ; and the large, round- 
quadrate Underleaves, connate with one leaf, with entire lateral 
margins and faintly four-lobed apices. (FiG. 37, nos. 1-5). 

DISTRIBUTION : Brazil: Rio de Janeiro, Gaudichaud (NY) . Ecua- 
dor: Cayambe, Jameson (NY). Venezuela: Valencia, Fendler, the 
type (H). 

REFERENCES: Stephani (/cone*, Mastigobryum no. 161). 



M. Fulford 106 Bazzania 

27. Bazzania Schwaneckiana (Hampe & Gottsche) Trevis. Mem. 
1st. Lomb. 13: 414. 1877. 

Mastigobryum Schwaneckianum Hampe & Gottsche, Linnaea 25 : 345. 1852. 

Plants scattered or growing in depressed mats, sometimes with- 
out leaves, mostly medium size, light green to olive-green, becoming 
brownish in the older portions: stems to 5 cm. long, with leaves 
mostly 1.5 mm. - 2.5 mm. wide, prostrate ; in longitudinal section 
the cells elongate, the medullary cells averaging 0.17 mm. long, the 
cortical shorter, both averaging 16/A in diameter, the end walls 
thin, the vertical walls uniformly thickened and containing frequent 
pits: the lateral branches mostly 5 mm. or more apart, diverging 
at a wide angle : flagelliform branches frequent, long, occasionally 
branched: rhizoids colorless, sparingly produced on the leaves of 
the flagelliform branches: the leaf -insertion curved in its upper 
half ; leaves approximate to imbricated, spreading, plane or nearly 
so, becoming deflexed on drying, unsymmetrically ovate, mostly 
1.2 mm. long, 0.56 mm. wide at the base, narrowing somewhat to 
the transversely truncate apex; the dorsal margin convex from a 
slightly rounded base, extending over one-half the stem, the ventral 
margin straight to slightly concave from a scarcely dilated base, 
the apex broad, rounded, entire to undulate or truncate, sometimes 
tending to become faintly two- to three-toothed, the margins entire 
to crenulate ; the leaf cells of the marginal one or two rows often 
forming a border, uniformly thick-walled, mostly 16/x X 10/i, the 
cells of the apical region 20//, - 24/i, thin-walled, the trigones very 
small, the median cells larger, those of the base to 50/* X 30/i, a 
vitta not differentiated, the cuticle verruculose: underleaves sub- 
imbricated, squarrose, attached in a straight line, connate by sev- 
eral cells with one leaf, quadrate to quadrate-orbicular, broader 
than the stem, mostly 0.34 mm. - 0.38 mm. long X 0.32 mm. - 0.38 
mm. wide, the apex undulate to two- or four-toothed or -lobed, the 
sinuses acute, narrow, the lateral margins lobed, convex from a 
straight base, the hyaline border complete, two to four cells wide, 
broadest at the apex, the cells similar to those of the chlorophyllose 
area but with thinner walls: leaves of the flagelliform branches 
ovate, mostly 0.16 mm. long, convex, the apices acute to bifid: 
sexual branches not seen. 

HABITAT : In depressed mats over logs and shaded banks. 

The species is best characterized by its medium size, the light 
green color, the rounded-entire or nearly entire leaf apices with 
crenulate margins, and the subquadrate, usually lobed underleaves 
with hyaline borders several cells wide, which are connate by a 
few cells with one leaf. (FiG. 38, nos. 1-14). 

The leaves while usually entire occasionally show a tendency to 
become three-toothed at the apex. These teeth are broad, obtuse, one 
or two cells high, with shallow, lunulate sinuses. The margins may 
be entire or more or less crenulate (FiG. 38, no. 6). The cells of the 
apical region are thin-walled, with small but distinct trigones, and 
the cell lumina are always rounded. The cells of the margin (one 
to several rows) are often longer and narrower, and have the walls 




FIG. 38. Bazzania Schwaneckiana (Hampe & Gottsche) Trevis. 
1. Dorsal view of portion of a stem, X 12. 2. Ventral view of portion of stem, 
X 30. 3. A leaf, X 30. 4. Leaf apices, X 30. 5. A cell from the apical 
portion of a leaf, X 400. 6. Cells from the apical portion of a leaf, X 260. 
7. Cells from the dorsal margin of a leaf, X 260. 8. Cells from the ventral 
margin of a leaf, X 260. 9. Cells from the basal portion of a leaf, X 260. 
10. Underleaf, X 30. 11. Underleaf and portion of a stem and leaf to show 
attachment, X 45. 12. Cells from the apical portion of an underleaf, the 
outer part hyaline, X 260. 13. Portion of a cross-section of a stem, X 260. 
14. Portion of a longitudinal section of a stem, X 260. Nos. 1, 3-14 drawn 
from material collected by EVANS 166 in Puerto Rico; no. 2 from a portion 
of the type material. 



M. Fulford 108 Bazzania 

uniformly thicker than those of the rest of the leaf, so that they 
form a border. This marginal border is best developed in plants 
growing in the more xerophytic situations. The cuticle is very 
rough. Large oil bodies were present in some of the plants col- 
lected by PAGaN, no. 274, in Puerto Rico. 

The vitta, which is characteristic of the leaves of many of the 
tropical species, is in B. Schwaneckiana not differentiated. While 
the basal portion of the leaf consists of a group of elongated cells 
arranged in rows, they are not sufficiently differentiated from the 
adjacent cells to form a distinctly differentiated area. The cell 
walls are thin and uniform (FiG. 38, nos. 5, 6, 7, 8 and 9) and the 
trigones are small but distinct. 

The underleaves are especially distinctive. They are subquad- 
rate in outline, broader than the stem, and are usually four-lobed 
at the apex (FiG. 38, nos. 2 and 10). The hyaline margin is two 
to four or five cells wide (FiG. 38, no. 12), continuous, and the cell 
pattern is the same as that of the chlorophyllose area. This hya- 
line border is often difficult to detect in the older underleaves where 
the chlorophyll content has disappeared or where the cell walls 
have become brown. It is always conspicuous in the younger 
leaves. The line of attachment on the stem is a straight line, in 
optical view the width of the stem, and in addition, a few cells of 
the ventral basal margin of one leaf. This latter character is not 
noticeable unless the plants are examined under high magnification 
(see FIG. 38, no. 11). It is always restricted to one side of the 
stem on a branch, and is especially conspicuous on that underleaf 
and leaf at the junction of the branch and stem. 

Many of the plants collected by EVANS and later PAGaN in 
Puerto Rico, and by Duss in Guadeloupe exhibit a tendency to shed 
their leaves, and many of the stems are without leaves throughout 
much of their length. This caducous habit seems to be restricted 
to the leaves only, for in no case were the underleaves absent even 
on those stems almost completely devoid of leaves. 

STEPHANI included this species under his subgenus Integrifolia 
because of the more or less entire character of its leaf apices. B. 
Schwaneckiana is without doubt very closely related to a small 
group of species restricted locally in the West Indies, namely B. 
Eggersiana, B. pycnophylla, B. armatastipula and B. cubensis. It 
shares with them size, habit of growth, color, texture, cell con- 
figuration and similarity of underleaves. All of them have the 
underleaf connate by a few cells with one leaf. In addition, all 
except B. cubensis have underleaves with conspicuous hyaline bor- 
ders. Because of this close natural relationship of the species with 
members of the Connatae it seems logical to transfer it to this 
group even though the leaf outline is not strongly serrate or the 
three teeth well defined. 

DISTRIBUTION: Dominica: Four Hand, Fishlock 12a (NY); without 
locality, Elliott, cited by Stephani (1908, 431). G ua del ou p e : without 
locality, 1'Herminier (H, Y, NY) ; Pointe Noire, Duss 43, 57 (NY). Mar- 



Bazzania 109 Connatae 

Unique : Calebasse, Morne Paillasse, Duss 38, 369 (NY). Duss 662 from 
Deux-Choux (NY), is a species of Frullania. Puerto Rico: without 
locality, Schwanecke, the type, 608 of Gottsche & Rabenhorst, Hep. eur. (Y, 
NY) ; near Adjuntas, E. G. Britton & Marble 2175 (NY) ; Luquillo Mountains, 
Heller 1158, 4653, 4654, 4717 (NY); El Yunque, Evans 166, 185 (Y, NY); 
Rio de Maricao, E. G. Britton 2503 (NY); without locality, Sintenis 12, cited 
by Stephani (1888a, 279); Orocovis, Pagan 22, 23 (C); Canovanas, Pagan 
274 (C). 

REFERENCES: Bescherelle (1893, 186); Stephani (1888a, 279; 1903, 22; 
1908, 431; Icones, Mastigobryum no. 22); Pagan (1939, 39). 

28. Bazzania pycnophylla (Taylor) Trevis. Mem. 1st. Lomb. 13: 

414. 1877. 

Mastigobryum pycnophyllum Taylor, London Jour. Bot. 5: 371. 1846. 

Plants growing in depressed mats, light green to olive-green, 
becoming tinged with brown ; stems slender, with leaves to 3 mm. 
broad, prostrate : the lateral branches 0.5 cm. or more apart, diverg- 
ing at a wide angle ; flagellif orm branches frequent ; rhizoids color- 
less, present on the bases of leaves of the flagellif orm branches: 
the leaf insertion little curved in the upper part ; leaves imbricated, 
spreading, becoming somewhat deflexed on drying, unsymmetri- 
cally ovate, to 1.4 mm. long, 0.55 mm. broad at the base, narrowing a 
little to the transversely truncate, serrulate to spinose, obscurely 
tridentate apex; the dorsal margin convex from a curved base 
which covers one-half the stem, the ventral margin straight to 
slightly concave, the base little dilated ; the apex broad, mostly ob- 
scurely tridentate, the teeth very short, acute, broad, the sinuses 
lunulate, the margins serrulate to spinose; the leaf cells subquad- 
rate to rectangular in outline, the cell lumina rounded, the cell 
walls of the apical portion and margins uniformly thickened, those 
of the interior thin with small but distinct trigones, cells of the 
apical portion 18/t X 18/x, of the dorsal lobe smaller, those of the 
base 36/i - 54/t X 27/* ; a vitta not differentiated ; the cuticle verru- 
culose: underleaves imbricated, a little broader than the stem, 
attached in a straight line, connate by several cells with one leaf, 
partly hyaline, quadrate, mostly 0.48 mm. X 0.48 mm., the lateral 
margins entire, serrate or dentate, the apex three- to six-toothed, 
the teeth two to five cells broad at the base, two to four cells high, the 
margins mostly serrate to dentate ; the cells of two sorts, those of 
the marginal rows (mostly of the teeth) hyaline, the walls uni- 
formly thin, verruculose, those of the interior chlorophyllose, the 
walls thicker, trigones small: leaves of the flagellif orm branches 
scale-like, ovate : male and female branches and sporophyte not seen. 

HABITAT : In depressed mats on logs and tree bases in woods. 

The distinguishing characteristics of the species are its pros- 
trate habit and olive-green color; the blunt, obscurely tridentate 
leaves which are serrulate to dentate in the apical portion ; and the 
quadrate underleaves, connate by a few cells to one leaf, with 
hyaline margins which are incised to form three to six, serrate to 
dentate teeth. (FlG. 39, nos. 1-10). 

The species is of the size and habit of the other tropical 
American Connatae. Its leaves are spreading and do not become 



M. Fulford 110 Bazzania 

deflexed on drying. The outstanding characteristic of the plant is 
its blunt, rarely obscurely tridentate leaves with conspicuously 
serrulate to spinose margins as shown in FIG. 39, nos. 2, 3, 4, 6. 

STEPHANI included the species among the Fissistipula but for 
obvious reasons it does not belong in that Section. 

B. pycnophylla is easily distinguished from B. Schwaneckiana 
because of its blunt, serrulate to spinose leaf apices. 

DISTRIBUTION: Jamaica : without locality, Swartz, the type (H, NY). 
Puerto Rico: Maricao, Pag&n no. 247*; Mt. Torrecillas, Bro. Hioram, 
12 (NY). Guadeloupe: without locality, rHerminier (H). 

REFERENCES: G. L. & N. (1847, 719); Lindenberg in Gottsche (1851, 73); 
Stephani (1908, 507; I cones, Mastigobryum no. 347). 

*I determined this specimen as B. Eggersiana for Dr. PAGAN, Bryol. 42: 
39. 1939, but until more material is available for study, it seems advisable to 
transfer it to B. pycnophylla because of the very short teeth on the leaves and 
the strongly serrate leaf apices. 

29. Bazzania Eggersiana (Steph.) Pagan, Bryol. 42: 39. 1939. 

Mastigobryum Eggersianum Stephani, Spec. Hep. 3: 468. 1908, 

Plants growing in depressed mats, medium size, light green to 
olive-green, a little pigmented with brown in the older portions: 
stems slender, to 5 cm. long, with leaves to 3 cm. broad, prostrate ; 
in longitudinal section the medullary cells averaging 0.17 mm. in 
length, the cortical shorter, both averaging 20/x in diameter, the 
end walls thin, the vertical walls thickened and containing frequent 
pits: lateral branches mostly 5 mm. apart, diverging at a wide 
angle; flagelliform branches numerous, long: rhizoids not seen: 
the leaf-insertion curved in its upper half; the leaves imbricated, 
spreading, plane or nearly so, sometimes a little deflexed on drying, 
unsymmetrical, ovate, mostly 1.4 mm. long, 0.65 mm. wide at the 
base, narrowing somewhat to the transversely truncate, tridentate 
apex; the dorsal margin convex from a curved base, extending 
over nearly one-half the stem, the ventral margin straight to 
slightly concave from a more or less dilated base, the apex broad, 
three-toothed, occasionally nearly rounded, the teeth mostly one 
to three cells long and two to four cells broad, the sinuses lunulate, 
the margins entire to serrulate, the leaf cells quadrate to rectangu- 
lar in outline, the cells of the apical region averaging 25/A X 25/x, 
those of the interior larger, those of the basal section 54/x X 30/x, 
not forming a vitta ; cell walls thin, the trigones small but distinct, 
the cell lumina rounded, the cuticle verruculose: underleaves im- 
bricated, a little squarrose in the upper part, attached in a straight 
line and connate by a few cells with one leaf, subquadrate, partly 
hyaline, as broad or broader than the stem, mostly 0.48 mm. - 0.56 



FIG. 39. Bazzania pycnophylla (Taylor) Trevis. 1. Portion of a 
plant, dorsal view, X 30. 2. Portion of a plant, ventral view, X 30. 3. Leaf, 
X 30. 4. Apices of leaves, X 30. 5. A cell from the apical portion of a leaf, 
X 350. 6-6a. Cells from the apical margin of a leaf, X 260. 7. Cells from 
the dorsal base of a leaf, X 260. 8. Cells from the basal portion of a leaf, 
X 260. 9. Underleaves, X 30. 10. Cells from the apical portion of an under- 
leaf showing the hyaline margin, X 260. Nos. 1-10 drawn from plants of the 
type material. 



M. Fulford 112 Bazzania 

mm. long X 0.56 mm. - 0.64 mm. broad, the apex with three to 
six, serrate to spinose-dentate teeth or lobes, the teeth to four cells 
broad, two to four cells high, the sinuses narrow, acute, the lateral 
margins serrate or with one to three teeth similar to those of the 
apex ; the cells of two sorts, those of the marginal one to four rows 
(mostly of the teeth), hyaline, strongly verruculose, with uniformly 
thickened cell-walls and no trigones, those of the interior chloro- 
phyllose, thin-walled, the trigones conspicuous, small : leaves of the 
flagelliform branches small, averaging 0.16 mm. long, the apex 
rounded, entire to crenulate: sexual branches not seen. 

HABITAT: On logs and bases of trees, woods. 

The species is best characterized by the dull green color; the 
transversely truncated leaf apices with mostly short, broad, entire 
to serrulate teeth; and the subquadrate underleaves with three to 
six, mostly hyaline, irregular, spinose-dentate teeth, three to six 
cells long, along the apical margins. (FiG. 40, nos. 1-10). 

The leaves, while not caducous are often brittle when dry and 
are easily broken, so that stems without leaves are not uncommon. 
The teeth show a wide degree of variation. There are usually 
three distinct teeth two to four cells broad and two to four cells 
high, but broad undulations with one of the cells forming the apex 
are frequent. The sinuses are mostly shallow and rounded. The 
margins are entire to serrulate, straight or occasionally undulate. 

The underleaves also, are variable. They are mostly subquad- 
rate in outline and as broad or broader than the stem. They are 
attached in a straight line and are connate by several cells to one 
leaf. The hyaline border is conspicuous and varies from one to 
three cells along the upper lateral margins and three to five cells 
at the apex. The three to six irregular lobes or teeth are usually 
entirely hyaline (see FIG. 40, nos. 2, 8, 9). The border is more 
conspicuous in young underleaves, mostly because in the older 
plants the walls of the hyaline cells become somewhat pigmented, 
while the chlorophyll content of the green cells of the interior tends 
to disappear, so that the strong contrast is lost. 

The color, size and habit of the plant, as well as its leaf apices 
and underleaves suggest a close relationship to B. Schwaneckiana. 
However, the teeth of the leaves are usually pronounced. The 
hyaline border of the underleaf of B. Schwwneckiana is continu- 
ous, and the apical portion undulate or lobed, with the margin 
entire, while in B. Eggersiana the hyaline border is restricted to 
the apex, and is toothed, and has a spinose-dentate margin. 

The species is distinguished from B. cubensis because of the 
hyaline margins of its underleaves, and from B. pycnophylla be- 

FiG. 40. Bazzania Eggersiana (Steph.) Pagan. 1. Portion of a plant, 
dorsal view, X 30. 2. Portion of a plant, ventral view, X 30. 3. Leaf apices, 
X 30. 4. Cells from an apical tooth of a leaf, X 260. 5. A cell from the 
apical portion of a leaf, X 350. 6. Cells from the dorsal margin of a leaf, 
X 260. 7. Cells from the ventral margin, including some from the basal 
portion of a leaf, X 260. 8. Underleaves, X 30. 9. Portion of the margin 
of an underleaf near the apex, X 260. 10. Portion of a transverse section 
of a stem, X 260. Nos. 1-10 drawn from a part of the type material. 



M. Fulford 114 Bazzania 

cause of the more pronounced teeth of its leaves, which have much 
less conspicuously serrulate or entire margins. STEPHANI included 
the species among the Grandistipula. This is the logical con- 
clusion if one is not too careful in the examination of the margins 
of the leaf apices, for on some of the stems many of the leaves 
show so few serrations that these are apt to be overlooked. On 
careful examination the connate habit of the underleaves also be- 
comes evident. This character, together with the cell pattern of the 
leaves, the general habit, and the leaf margins, especially those with 
many serrations, point to a closer relationship to B. pycnophytta 
and B. armatastipula, than to any member of the GrandistijnUae. 

DISTRIBUTION : C u b a : Final de Santa Anna, 2400', Eggers, the type (H) ; 
Monte de la Prenda, 2400', Eggers, 5191c (NY) ; Banao Mountain, Brother 
Le*on & Per. Roca, 8340 (NY). The specimen collected by PAGAN 247 at 
Maricao, Puerto Rico, should be referred to B. pycnophylla. 

REFERENCES: Stephani (Icanea, Mastigobryum no. 160). 

30. Bazzania armatistipula (Steph.) comb. nov. 

Mastigobryum armatistipulum Stephani, Spec. Hep. 3: 490. 1908. 

Plants growing m depressed mats, medium size, light green to 
olive-green, becoming pigmented with brown in the older portions : 
stems slender, to 5 cm. long, with leaves to 3 mm. broad, prostrate ; 
in longitudinal section the cells elongate, the medullary cells av- 
eraging 0.17 mm. long, the cortical shorter, both averaging 20/x in 
diameter, the end walls thin, the vertical walls thickened and con- 
taining frequent pits; lateral branches 5 mm. or more apart, 
diverging at a wide angle ; the flagellif orm branches frequent, long ; 
rhizoids not seen : the leaf insertion curved in its upper part ; the 
leaves approximate to imbricated, spreading, plane or nearly so, 
unsymmetrical, ovate, mostly 1.4 mm. long, 0.65 mm. wide at the 
base, narrowing somewhat to the transversely truncate, tridentate 
apex ; the dorsal margin convex from a curved base, extending over 
nearly one-half the stem, the ventral margin straight to slightly 
concave from a more or less dilated base ; the apex broad, the teeth 
acute, one to five cells broad at the base, two to five cells long, the 
sinuses lunulate, the margins serrate; the leaf cells quadrate to 
rectangular in outline, the cells of the apical region averaging 
25/x X 25/x, those of the interior larger, those of the basal portion 
54/A X 36/A, not forming a vitta, the walls of the marginal rows 
more or less uniformly thickened, those of the interior cells thin 
with small but distinct trigones, the cell lumina rounded, the cuticle 
verruculose: underleaves approximate to imbricated, attached in 
a straight line, connate by a few cells with one leaf, subquadrate, 
mostly 0.56 mm. X 0.56 mm. - 0.64 mm. broad, with a hyaline 
border, the lateral margins bulging, dentate to serrate, the apex of 
three to six spinose-dentate teeth or lobes, the teeth to four cells 
broad, two to four cells high ; the hyaline border one to five cells 
broad, the cells averaging 18/A X 12/x; the chlprophyllose cells 
averaging 20/i, the trigones conspicuous, the cuticle verruculose: 
leaves of the flagelliform branches scale-like, ovate, the apex mostly 
rounded : sexual branches not seen. 



LJ 



rvr\vJL^s<v)\ " 4 /^vrN^nV 




FIG. 41. Bazzania armatistipula (Steph.) Fulford. 1. Portion of a 
stem, ventral view, X 30. 2. Portion of a stem, dorsal view, X 30. 3. Leaf, 
X 30. 4. Leaf apices, X 30. 5-5a. Apical teeth, X 260. 6. A cell from the 
apical portion of a leaf, X 350. 7. Cells from the dorsal margin of a leaf, 
X 260. 8. Cells from the basal portion of a leaf, X 260. 9. Underleaf, X 30. 
10. Portion of the apical region of an underleaf showing the hyaline border, 
X 260. 11. Underleaf and portion of stem and a leaf to show line of at- 
tachment, X 30. Nos. 1-11 drawn from a portion of the type material. 



M. Fulford 116 Bazzania 

HABITAT : In depressed mats over logs and tree bases in woods. 

The distinguishing characteristics of the species are its light 
to olive-green color; the spreading, tridentate leaves with con- 
spicuously serrulate to dentate apical margins; and the variously 
spinose-serrate, toothed underleaves which are bordered by several 
rows of hyaline cells in the apical regions. (FiG. 41, nos. 1-11). 

The species has much the same habit as the other members of 
the Connatae just described. It can, nevertheless, readily be dis- 
tinguished from them because of its strongly three-toothed leaf 
apices which have conspicuously serrulate to spinose margins as is 
shown in FIG. 41, nos. 1, 4, 5. The leaf apices of B. cubensis some- 
times are very similar but the underleaves are always chlorophyllose 
throughout so that the two need never be confused. 

The underleaves are similar to those of B. pycnophylla and B. 
Eggersiana, and cannot be used as a diagnostic characteristic dis- 
tinguishing these species from one another. They are always 
connate by a few cells with the ventral base of one leaf as is shown 
in FIG. 41, no. 11. 

STEPHANI included M. armatastipulum as the only American 
species of the Serrulata but did not consider it among the connate 
group. 

DISTRIBUTION : C u b a : Sierra Maestro, Bro. Leon, Clement, and M. Roca, 
10471 (NY); Banao Mountains, Bro. Leon and M. Roca, 8075 (Y, NY). (?). 
Jamaica : without locality, Borgensen, the type (H) ; near Hardware 
Gap, 4000 ft, Underwood 2242 (Y, NY); John Crow Peak, 5500-5800 ft, 
Underwood 855 (NY); Blue Mountains, Elizabeth Britton 1193 (NY); 
Chapelton to Bull Head, Underwood 3400 (NY) ; without locality, Wilson 186 
(NY). Puerto Rico: Monte Alegdllo, 500-900 m., Elizabeth Britton, 
2650 (Y, NY); Maricao, Britton and Cowell 4251 (NY). 

REFERENCES: Stephani (/cones, Mastigobryum no. 262). 

31. Bazzania cubensis (Gottsche) Pagan, Bryologist 42: 38. 1939. 

Mastigobryum cubense Gottsche, in Stephani, Hedwigia 24: 248. pi. 3, fig. 1. 
1885. 

Plants in depressed mats, medium size, olive-green tinged with 
brown : stems slender, to 6 cm. long, with leaves to 3.5 mm. broad, 
prostrate ; in longitudinal section the cells elongate, the medullary 
cells averaging 0.17 mm. long, the cortical cells shorter, both 
averaging 20/x in diameter, the vertical walls uniformly thickened 
and containing frequent pits, the end walls thin; lateral branches 
mostly 1 cm. or more apart, diverging at a wide angle ; flagellif orm 
branches frequent, long; rhizoids colorless, present on the leaves 

FIG. 42. Bazzania cubensis (Gottsche) Pagan. 1. Portion of a stem, 
dorsal view, X 30. 2. Portion of a stem, ventral view, X 30. 3. A leaf, X 30. 
4. Leaf apices, X 30. 5. A cell from the apical portion of a leaf, X 350. 
6. Cells from an apical tooth of a leaf, X 260. 7. Cells from the dorsal 
margin of a leaf, X 260. 8. Cells from the basal portion of a leaf, X 260. 
9. Underleaves, X 30. 10. Cells from the apical portions of two underleaves, 
X 260. 11. Underleaf and portion of a stem and leaf to show attachment, 
X 30. Nos. 1-11 drawn from a portion of the type material. 



M. Fulford 118 Bazzania 

of flagelliform branches; leaf insertion little curved in the upper 
part; leaves imbricated, spreading, a little convex, deflexed when 
dry, unsymmetrically ovate, 0.9 mm. - 1.6 mm. long, 0.6 mm. - 0.7 
mm. broad at the base, narrowed a little to the mostly transversely 
truncate, tridentate apex ; the dorsal margin convex from a curved 
base which covers one-half the stem, the ventral margin straight 
or slightly concave, the base scarcely dilated; the apex broad, the 
teeth acute, large to obscure, two to eight cells long and three to 
six cells broad at the base, the sinuses lunulate, the margins mostly 
slightly serrulate ; the leaf cells thin- walled, the cell lumina angular- 
rounded, the trigones conspicuous, small, rarely coalesced, the cells 
of the apical portion 22^ - 25^ X 25/x - 30/x, the marginal row 
smaller and with thicker walls, the cells of the median portion 
larger, those of the basal area 36/x - 54/x X 27/x ; a vitta not differ- 
entiated; the cuticle very strongly verruculose; underleaves quad- 
rate, little broader than the stem, imbricated, attached in a straight 
line, narrowly connate with one leaf, squarrose in the upper part, 
the lateral margins straight or slightly convex, serrate and lobed, 
the apex toothed, the teeth irregular, two to five cells broad at the 
base, two to four cells high, the margins entire to serrate; the 
cells 25ju, X 25/x, becoming longer in the central portion, the cell 
walls thin, the trigones conspicuous, often becoming coalesced, the 
cell lumina angular-rounded, the cuticle strongly verruculose : male 
and female branches and sporophytes not seen. 

HABITAT: On logs and bases of trees, woods. 

The distinguishing characteristics of the species are its olive- 
green to brown color; the deflexed leaves, with irregularly tri- 
dentate, mostly serrate apices, and large thin-walled cells with 
conspicuous trigones ; and the quadrate underleaves, connate by a 
few cells with one leaf, coarsely toothed and incised at the apex and 
often along the margins. There are no hyaline cells in the under- 
leaf. (Fio. 42, nos. 1-11). 

The plants are approximately the same size as the other tropical 
American members of this Section, but they differ in appearance, 
in that the leaves are always strongly deflexed when dry, and the 
green color is always tinged with brown, even at the growing tip, 
so that the plants are olive-green or darker brown. 

The leaves are usually tridentate, with the teeth large, usually 
broadly triangular. However, on a single stem it is not unusual 
to find all sorts of variations in the size and shape of the teeth, 
from the large sort mentioned above, to small ones of one or two 
cells (see FIG, 42, nos. 2, 3, 4). The margins are serrate to a 
greater or lesser degree. The marginal cells have thick walls (see 
FIG. 42, no. 6), and the cuticle is conspicuously verruculose to 
rough papillose. 

The underleaves are very similar to those of the other American 
species of the Connatae except that all of the cells are chlorophyllose. 
They are attached in a straight line to the stem and to several cells 
of the ventral base of one leaf (see FIG. 42, no. 11). As in the 
case of B. Schwaneckiana this attachment to the leaves is not 
noticeable except under high magnification. 



Bazzania 119 Connatac 

STEPHANI considered M. cubense to be a member of the Fissi- 
stipula Section. The species certainly does not belong to that 
group since the underleaves are not deeply incised (see FIG. 42, 
no. 9). He likewise did not recognize the fact that the leaves are 
serrate, some of them strongly so, and that an underleaf is connate 
with one leaf for a short distance (see FIG. 42, nos. 2, 4, 11). 

DISTRIBUTION : Cuba: Mt. Verde, Wright, Hep. Cub. Wrightianae, the 
type (Y, NY) ; Sierra Nipe, near Woodfred, Oriente, 450-550 m., Shafer 3344, 
3434 (NY); Sierra Maestra, Bro. Clement 343 (NY). J am ai c a : sum- 
mit Bull Head, Underwood 3390 (NY). Puerto Rico: Maricao, 
Pagan 216a, 237, 251 (F). 

REFERENCES: Stephani (1908, 507; /cones, Mastigobryum no. 344). 

32. Bazzania peruviana (Lehm. & Lindenb.) Trevis. Mem. 1st. 

Lomb. 13: 414. 1877. 

Jungermannia peruviana Lehman & Lindenberg, in Lehman, Pug. PL 5: 18. 

1833. 

Mastigobryum peruvianum Nees in G. L. & N., Syn. Hep. 220. 1845. 
Mastigobryum peruvianum var. ft minimum Gottsche & Schiffner, in Schiffner, 

Lebermoose. Forschungsreise S.M.S. "Gazelle" IV, 17. pi. 4, fig. 17-18. 

1888. 
Mastigobryum Lechleri Stephani, Hedwigia 25 : 134. pi. 6, fig. 10-14. 1886. 

Plants medium size, in deep tufts or mats, dark green, becom- 
ing pigmented with brown in the older portions: stems 5 cm. or 
more in length, with leaves to 3.5 mm. broad, prostrate ; stem cells 
in longitudinal section to 0.17 mm. long, the cortical shorter, av- 
eraging 20/x in diameter, the vertical walls uniformly thickened 
and containing frequent pits, the end walls thin; lateral branches 
frequent, mostly 5 mm. apart, diverging at an acute angle : flagelli- 
form branches frequent; rhizoids rare, colorless, on the bases of 
the leaves of the flagelliform branches; the line of leaf insertion 
little curved in the upper part, the leaves imbricated, plane, be- 
coming a little deflexed on drying, unsymmetrically ovate, straight, 
1.5 mm. - 2 mm. long, 1 mm. broad at the base, narrowed a little 
to the transversely truncate, sharply tridentate apex; the dorsal 
margin arched from a curved base, covering up to one-half the 
stem, the ventral margin somewhat concave, often conspicuously 
dilated, entire, the apex three-toothed, the teeth variable, mostly 
sharply acute, three to eight cells long, two to four cells broad, the 
sinuses mostly deeply lunulate, the margins serrulate to dentate; 
leaf cells thin-walled, the cell lumina rounded, trigones small, the 
cuticle verruculose; cells of the apical region averaging 20/x - 24/x, 
of the median portion larger, and of the base to 56/t X 24/A, a vitta 
not differentiated : underleaves imbricated, subquadrate in outline, 
6 mm. - 8 mm. long and broad, broader than the stem, attached in 
a straight line, conspicuously connate with a pair of leaves, be- 
coming squarrose and strongly recurved in the upper half, the 
apex more or less lobed, the lobes serrulate to spinose-dentate, the 
lateral margins convex, entire; the hyaline border four to eight 
cells broad, thin-walled, the chlorophyllose cells with thicker walls 




FIG. 43. Bazzania peruviana (Lehm. & Lindenb.) Trevis. 1. Portion 
of a plant, ventral view, X 15. 2. Portion of a stem and leaf, dorsal view, 
X 30. 3. Apices of leaves, X 30. 4. Apical tooth of a leaf, X 310. 5. A cell 
from the apical portion of a leaf, X 400. 6. Underleaves, X 30. 7. A portion 
of an underleaf to show hyaline border, X 310. 8. Portion of a stem, ventral 
view, X 15. 9. A cell from the apical portion of a leaf, X 400, Nos. 1-7 
drawn from the type of J. peruvicwia; 8-9 from the type of M. Lechleri from 
Chile. 



Bazzania 121 Connatae 

and small trigones: leaves of the flagelliform branches scale-like, 
ovate, the margins serrulate to dentate: sexual branches and 
perianths not seen. 

HABITAT: In depressed mats or ascending tufts on soil, rocks, 
and tree bases. 

The distinguishing characteristics of the species are its medium 
size, the dark green or yellowish color; the sharply three-toothed 
leaves which are entire or serrate along the apical margin; and 
the connate (with a pair of leaves) underleaves which are 
squarrose-recurved in the upper half, and have hyaline borders 
two to eight cells broad, with serrate to spinose-dentate apical 
margins. (FlG. 43, nos. 1-9). 

The species exhibits much variation in its leaves and under- 
leaves. On most well developed plants the teeth are long, narrowly 
acute to acuminate, and usually end in a point of several cells. On 
less well developed plants this characteristic is less often observed, 
for most of the teeth are short, acute, with only an occasional 
tooth long-pointed. The margins of the teeth also vary consider- 
ably. In the well developed plants with long teeth the margins are 
crenulate to serrate, often strongly so. However, the amount and 
degree of serration is not constant on all the leaves of a single 
plant, nor on the leaves of the various plants of a mat (see FIG. 
43, nos. 1, 3 and 4). The serrations tend to become fewer and 
smaller on the less well developed plants and are often entirely 
absent on many of the leaves (see FIG. 43, no. 8). The thickness 
of the cell walls, the size of the trigones, and the roughness of the 
cuticle show the usual range of variation. 

The underleaves are always conspicuously connate with a pair 
of leaves as seen in FIG. 43, no. 1. The connecting bands are broad 
and can easily be determined. The upper portions of the under- 
leaves are strongly recurved on well developed plants, but tend to 
become less so in the poorer forms as is seen when comparing FlG. 
43, nos. 1 and 8. The hyaline margin is usually very conspicuous 
and may be as much as ten cells broad in the apical part, but here 
again one finds much variation, for on some plants only two or 
three rows of cells are hyaline (see FIG. 43, nos. 1, 6, 7 and 8). 
On plants with underleaves having this narrow border the under- 
leaf just below a flagelliform branch usually is more typical, i.e. 
the hyaline border is very wide. The apical portion is more or less 
four-lobed and the margin is serrate and dentate. 

Plants having leaves with very rough cuticle, short teeth with 
distantly serrate margins, and underleaves with narrow hyaline 
borders, occur more frequently in the southern part of Chile than 
does the form described above, although there is no complete 
geographical segregation of the forms. The plants collected by 
SKOTTSBERG on the Swedish Pacific Expedition, 1916-1917, were 
for the most part robust, the teeth of the leaves were long on some 
plants, short on others, the hyaline border of the underleaves 



M. Fulford 122 Bazzania 

varied in width as described above, and the cuticle was always 
more verruculose than that of the type specimen, often becoming 
very roughly warty. 

The var. minimum of GOTTSCHE and SCHIPPNER has the char- 
acteristics of the species except for its smaller size and much 
smaller underleaves. I believe that it should be considered as a 
habitat variation rather than a variety. It appears to be a plant 
which grew in a very moist, perhaps densely shaded habitat. 

STEPHANI originally described the underleaves of his M. Lechleri 
as being free, but in the Icones, Mastigobryum no. 167, has shown 
several of the underleaves as connate with single leaves some 
on the left and some on the right side of the stem. Since plants 
with underleaves connate with one leaf always show this char- 
acteristic restricted to the same side of the stem for all of the 
leaves on a branch, it is obvious that the underleaves of STEPHANI'S 
plant were connate with pairs of leaves. M. Lechleri is a rather 
small form of B. peruviana. The portion of the type studied had 
the underleaves connate with pairs of leaves but the line of attach- 
ment was broader for one leaf than for the other of a pair as 
indicated in FIG. 43, no. 8. 

The species is closely related to B. Skottsbergii which follows. 

DISTRIBUTION : Peru: without locality or collector's name, Hb. Gottsche, 
the type (H) ; without locality, Jameson (NY). Chile : Valdivia, Hahn 
(H); Valdivia, Lechler, the type of M. Lechleri (H) ; Corral, Thaxter 142, 
149 (Y, H); Curico, Wollemeyer 279a (H) ; without locality, H. C. Lorenz 
(NY); Port Montt, Bro. Claude-Joseph 3216 (Y) ; Chiloe* Island, Anderson 
(NY); also Cunningham 1445 (NY); Chonos Archipellago, Darwin Bay, 
without collector's name 442 (NY) ; Panguipulli, Padre Hollermeyer 279 (Y); 
also Bro. Claude-Joseph 2322, 2322a (Y) ; Str. Magellan, Port Otway, Cun- 
ningham 235 (NY); Gray Harbor, Cunningham 147 (NY); Hall Bay, Cun- 
ningham 212 (NY) ; South Channel, without collector's name (as M. cerinum) 
(H) ; Tuesday Bay, Naumaun, the type of the var. 13 minimum (H), also nos. 
68, 133 p.p. (H); Desolation Island, Duse"n cited by Stephani (1901, 22). 
Patagonia : "occidentalis in insula Newton in Terra", cited by Stephani 
(1900, 51); without locality, Hatcher, cited by Evans (1903, 42). Juan 
Fernandez Islands: Skottsberg (Swedish Pac. Exp. 1916-1917), Baz- 
zania nos. 1, 2, 7, 10, 14 (S). 

REFERENCES: Evans (1903, 42); G. L. & N. (1844-1847, 215); Lindenberg 
& Gottsche (1851, 25. pi. 6. fig. 1-6); Stephani (1900, 51; 1901, 22; 1908, 457, 
469; 1911, Icones, Mastigobryum nos. 114, 167). 

33. Bazzania Skottsbergii (Steph.) comb. nov. 

Mastigobryum Skottsbergii Stephani, in Skottsberg, K. Svensk. Handl. 46: no. 
9. 60. fig. 22, i, k. 1911. 

This species is very similar to B. peruviana in color, size, growth 
habit, etc. It is distinct, however, in that the underleaves are 
chlorophyllose throughout or have only a single row or only 
scattered hyaline cells along the margins. The margin is usually 
distantly or obscurely serrate or may even be entire. The under- 
leaves of B. peruviana are bordered by three to six rows of hyaline 
cells and the margin is conspicuously serrate and may be dentate. 



Bazzania 123 Connatae 

The apices of the leaves are usually less serrate than those of 
B. peruviana. The cells are slightly larger and the trigones more 
pronounced. The cuticle is faintly to strongly verruculose. 

HABITAT : Over rocks and on bases of trees. 

DISTRIBUTION : Juan Fernandez: Masatierra, Skottsberg (Exp. 
Suec. 1907-09), the type (L); various localities (Exp. Suec, Pac. 1916-17), 
Skottsberg 3, 5, 6, 8, 9, 12, 13 (S). 

ChiloS, western Patagonia, southern Patagonia and the Falkland Islands 
are also cited by Stephani. 

REFERENCE: Stephani (1924, 480; /cones, Mastigobryum no. 445). 

The species seems to be very close to or identical with M. cerinum St. col- 
lected by DUSEN in western Patagonia. The type material of this latter 
species has not been available for study so that detailed statements con- 
cerning the species cannot be made at this time. However, there are plants 
in the herbaria of the New York Botanical Garden and Yale and Harvard 
Universities which were collected in Patagonia by DUSEN and determined by 
STEPHANI as Jd cerinum. 

These plants also are similar to B. peruviana except for several details. 
The leaf cells are slightly larger, the walls thicker, the trigones more pro- 
nounced and the cuticle more often is conspicuously warty. The principle 
distinction is in the underleaves. They are as large as the underleaves of 
B. peruviana but the number of hyaline cells is limited to a marginal row, 
or scattered individuals across the top of the underleaf. The cell walls are 
similar to those of the leaf, the trigones are conspicuous, and the cuticle is 
strongly verruculose. The apical portion is more or less four-lobed and the 
margins are serrate to dentate, but a little less so than in B. peruviana. These 
plants are very similar to (if not identical with) the type material of B. 
SkottsbergiL 

The following specimens have been examined: Patagonia, San Pedro 
Island, Dus^n (H) ; Desolation Island, Tierra del Fuego, Duse"n 143 (Y, NY). 

34. Bazzania novae-zelandiae (Mitten) Bescherelle & Massalonga, 

in Mission Sci. du Cap Horn 1882-1883. V. Bot. 

233. 1889, by P. HARIOT and others. 

Mastigobryum novae-zelandiae Mitten, in Hooker, Bot. Ant. Voy. 2': Fl. N. 
Zel. 148. pi. C, fig. 6. 1854. 

The plants of the type material collected on North Island, New 
Zealand, are robust, to 8 cm. long and with leaves to 5 cm. broad. 
The teeth are long, acute, and sometimes acuminate but the margins 
are only rarely faintly serrulate. The underleaves are very large, 
with broad hyaline borders. The margins are undulate and entire 
or nearly so. They differ from B. peruviana in that in the latter 
the underleaves are serrulate-dentate, particularly along the more 
or less four-lobed apices. 

B. novae-zelandiae has been recorded from Patagonia and the Straits of 
Magellan by BESCHERELLE and MASSALONGA, and from western Patagonia, 
(collected by DUSEN) by Stephani. None of these specimens has been 
examined. 



Section 3. Fissistipulae 

The Section is characterized by having the three-toothed leaves 
not serrate along the margins, and without appendages or con- 
spicuous auricles at the ventral bases; the underleaves are deeply 
incised (approximately one-half or more) or long-toothed. 

Unfortunately none of the American species which STBPHANI 
included in this Section actually possess a combination of characters 
which would include them in this subdivision. None of the species 
have deeply incised or long-toothed underleaves. 

With the exception of M. heterophyllum and M. Burchellii the 
species have been transferred to sections with which they are more 
closely related and are discussed under those sections. 

It has not been possible to examine authentic material of either 
M. Burchellii or M. heterophyllum. The figures in the Icones, 
Mastigobryum nos. 340, 346 and 442 indicate that these species 
also should be transferred, possibly to the Grandistipulae Section. 



Section 4. Appendiculatae 

Plants with leaves subentire to sharply three-toothed ; attached 
in a hook-formed line; the ventral bases dilated to a greater or 
lesser degree to form entire, serrulate, or appendiculate auricles; 
leaf cells thin- walled, the trigones large; the underleaves large, 
cordate, attached in a recurved line. 

Key to the Species 

1. Plants very large, the leaves subentire . . . . 42. B. canelensis (p. 152) 
1. Plants medium to large, the leaves three-toothed. 

2. Ventral auricles large, undulate, the margins serrate to dentate, the 
margins of the underleaves similar; cuticle very rough, 

35. B. asperistipula (p. 125) 
2. Ventral auricles and margins of the underleaves not as above. 

3. Plants very large; underleaves with one or more very long, sharp 
teeth at right angles to the lateral margins, 

41. B. acanthoetipa (p. 151) 

3. Plants medium size to large; underleaves with shorter teeth, 
lobed or with entire margins. 

4. Underleaves deeply four-toothed or -lobed, often with teeth 
several cells long on the margins; ventral auricles of the 

leaves well developed 40. B. teretiuscula (p. 144) 

4. Underleaves never deeply four-lobed; the ventral auricles of 
the leaves large or small. 

5. Margins of the underleaves often angular and incised; 
cell cavities stellate, trigones very large, rounded; leaf- 
lets of the flagelliform branches large, 

36. B. falcata (p. 129) 

5. Margins of the underleaves usually entire, undulate; cell 
cavities angular-rounded to stellate, trigones large to 
medium size, often rounded. 

6. Leaves not conspicuously narrowed at the apex; 
ventral appendages absent or poorly developed; the 
teeth broad, equilateral, the cells averaging 16j - 
20/4, thin-walled, the cuticle verruculose; leaflets of 
the flagelliform branches large, 

37. B. Hookeri (p. 135) 

6. Leaves narrowed at the apex, the teeth narrow, 
sharply pointed; the ventral auricles conspicuous, 
undulate, entire or with appendages. 
7. Leaves spreading, the cells averaging 18/& - 20/i 
in diameter; leaflets of the flagelliform branches 
scale-like ... 39. B. Liebmanniana (p. 142) 
7. Leaves falcate, the cells averaging 24/t in di- 
ameter; leaflets of the flagelliform branches 
large 38. B. robusta (p. 140) 

35. Bazzania asperistipula (Steph.) comb. nov. 

Mastigobryum aspertetipulum Stephani, Spec. Hep. 6 : 453. 1924. 

Plants in deep erect tufts, very large, light yellow-green, be- 
coming light brown in the older portions: stems stout, 10 cm. or 










Fio. 44. Bazzania asperistipula (Steph.) Fulford. 1. Portion of a 
plant, dorsal view, X 30. 2. Portion of a plant, ventral view, X 10. 3. Leaf 
apices, X 30. 4. A cell from the apical portion of a leaf, X 350. 5. Cells of 
an apical tooth, X 260. 6. Cells from the dorsal base of a leaf, X 260. 
7. Underleaf, X 30. 8. Basal portion of an underleaf, X 30. 9. Cells from 
the apical portion of an underleaf, X 260. 10. Ventral appendages of two 
leaves, X 60. Nos. 1-10 drawn from a portion of the type material. 



Bazzania 127 Appendiculatae 

more long, with leaves to 6 mm. broad, erect; stem cells in longi- 
tudinal section elongate, averaging 0.18 mm. long, the cortical 
shorter, both averaging 18/4, in diameter, the vertical walls strongly 
thickened and containing frequent pits, the end walls thin, the 
walls of the cortical and subcortical layers strongly pigmented with 
brown, the medulla hyaline : lateral branches infrequent, 1 cm. or 
more apart, diverging at a wide angle; flagelliform branches fre- 
quent, long ; rhizoids colorless present on some of the leaves of the 
flagelliform branches : the leaf insertion curved in its upper part, 
the dorsal end turned downward forming a hook; leaves always 
spreading, subimbricated to imbricated, unsymmetrically long- 
ovate, a little falcate, to 4 mm. long, 2 mm. broad at the base, 
narrowing to the rather broad, transversely truncate, tridentate 
apex; the dorsal margin strongly curved from a deeply cordate 
base, extending across the stem and somewhat beyond, the ventral 
margin concave, the base conspicuously auricled, the dilation ob- 
long, undulate and lobed, crenulate and serrate ; the apex strongly 
three-toothed, the teeth broadly triangular, acute, five to ten cells 
long and five to ten cells broad at the base, the sinuses deep, 
broadly acute, the margins straight to repand ; the leaf cells thin- 
walled, the cell lumina angular-rounded, the trigones small, con- 
spicuous, often becoming confluent, the cuticle strongly verruculose ; 
cells of the apical region and dorsal lobe quadrate in outline, 
averaging 22/i X 22/i, those of the marginal row and ventral auricle 
somewhat smaller, cells of the median portion longer, 36/A X 22u, 
and those of the basal portion 45/t X 27/*, a vitta not differentiated : 
underleaves imbricated, twice as broad as the stem, subquadrate 
in outline, attached in a recurved line, somewhat squarrose, the 
margin repand, 0.8 mm. - 1.4 mm. long and wide, the base strongly 
cordate, the auricles large, undulate, the margins lobed, serrate and 
often short ciliate, the apex and lateral margins lobed and toothed, 
the lobes broad and shallow, the sinuses lunulate, shallow, the 
margin crenulate to serrate, often dentate or short ciliate, the cells 
similar to those of tha leaf, those of the marginal row averaging 
22/x X 22/1, those of the basal portion larger, the trigones as in the 
leaf; leaves of the flagelliform branches to 0.35 mm. long, ovate, 
the apex acute or bifid with long, one-celled teeth : female branches 
occasional, solitary, one to several on a stem, the bracts and brac- 
teoles similar, little pigmented with brown; the outermost series 
ovate, bifid to trifid; the intermediate series long, broadly ovate, 
the margins entire to crenulate or dentate, one-third to one-fourth 
divided into two to four crenulate laciniae, the cells uniform 
throughout, 90/x X 20/i, thin-walled; the innermost series broadly 
ovate (immature), one-third to one-fourth divided into usually 
three, long, dentate to ciliate laciniae, the cells uniform, elongate, 
72/i - 90/4 X 20/i, the cell wall thin: perianth (immature) mouth 
long laciniate, the laciniae crenulate to ciliate, mostly twelve or 
more cells long: male branches and sporophytes not seen. 

HABITAT: In deep tufts on stumps and logs. 

The species is best characterized by its very large size, the long, 
seldom branched stems, the light yellow-green color; the subim- 
bricated, elongate, tridentate leaves with large, serrate to dentate, 
lobed auricles, and thin-walled leaf cells with conspicuous trigones 




FIG. 45 Bazzania asperiatipula (Steph.) Pulford. 1. Portion of the 
stem, longitudinal section, X 260. 2. Portion of stem, transverse section, 
X 260. 3. Female bract of intermediate series, X 30. 4. Female bract of 
innermost series (immature), X 30. 5. Portion of one of the laciniae of a 
bract of this series, X 100. 6. One of the laciniae of the mouth of the perianth, 
X 100. Drawn from the type material. 



Bazzania 129 Appendiculatae 

and verruculose cuticle; and the subquadrate underleaves with 
strongly cordate bases, and undulate, lobed, crenulate to dentate, 
short ciliate margins. (FIGURES 44, nos. 1-10; 45, nos. 1-6). 

The material available for study did not show much variation 
in the plants. The leaves are always light green in color, strongly 
three-toothed, and have the ventral appendages well developed. 
The appendage is fairly large, oblong, and is not sharply delimited 
from the rest of the leaf. The margin is lobed, crenulate and dis- 
tantly serrate to dentate as shown in FIG. 44, no. 10. The under- 
leaves also show little variation. They are all strongly cordate and 
have a variously lobed or sometimes toothed margin. These lobes 
and teeth are usually crenulate to serrate or occasionally short 
ciliate. The cilia are usually found on the basal portion. The basal 
auricles are large and may overlap (see FIG. 44, nos. 2, 7 and 8). 

STEPHANI'S unpublished drawings (Icones, Mastigobryum nos. 
307 and 373), made from plants of the type material collected by 
TURCKHEIM in Guatemala show two sorts of underleaves. The one 
has a uniformly serrate margin, while the other is broadly lobed 
or has broad, shallow teeth. 

DISTRIBUTION : Guatemala: Alta Verapaz, Coban, 1600 m., Turck- 
heim 5817, the type (NY); the same, 5503 as M. teretiusculum (NY); the 
same, 5582 (H). Costa Rica: Coliblanco, 1950 m., Maxon 237 (NY); 
Cerro de La Carpintera, Standley 35632 (W) ; Oroei, Standley 39650 (W); 
Alto de La Estrella, Standley 39106, 39403 (W) ; El Muneco, Standley 51338 
(W). Colombia: without locality, Wallace (NY). Venezuela: 
Caracas, without collector's name (H). 

REFERENCES: Stephani (Icones, Mastigobryum nos. 307, 373); Herzog 
(1938, 19). 

36. Bazzania falcata (Lindenb.) Trevis. Mem. 1st. Lomb. 13: 415. 

1877. 

Mastigobryum falcatum Lindenberg, in G. L. & N., Syn. Hep. 231. 1845. 
Mastigobryum tocutianum Gottsche, in Stephani, Hedwigia 25: 236. pi. 2. fig. 

18-20. 1886. 
Bazzania ancistrodes Spruce, Trans. & Proc. Bot. Soc. [Edinburgh] 15: 380. 

1885. 

Mastigobryum ancistrodes Stephani, Spec. Hep. 3: 501. 1908. 
Mastigobryum armatum Stephani, Spec. Hep. 3 : 528. 1909. 

Plants large, ochraceous yellow to dark brown, greenish in the 
younger portions: stems robust, to 10 cm. long, with leaves to 
6 mm. broad, depressed to ascending; in cross section the cells 
thick-walled, averaging 25/x in diameter ; lateral branches frequent, 
diverging at a wide angle; flagelliform branches frequent, short: 
rhizoids colorless, in tufts, from the bases of the leaves of flagelli- 
form branches : the line of leaf insertion curved in its upper half, 
the dorsal end turned downward forming a hook; leaves densely 
imbricated, convex, deflexed, connivent when dry, unsymmetrically 
long-ovate, somewhat falcate, to 3.5 mm. long, 2 mm. broad at the 
base, narrowing to the mostly 0.65 mm. broad, truncate, tridentate 
apex ; the dorsal edge strongly convex from a deeply cordate base, 
extending across the stem and somewhat beyond, the ventral edge 



M. Fulford 130 Bazzania 

concave, the base conspicuously auricled, the dilation large, oblong 
to rounded, undulate, entire to toothed ; the apex three-toothed, the 
teeth broadly triangular, acute, five to eight cells broad at the base, 
to six cells high, widely spreading, the sinuses broad, lunulate, the 
margins straight to repand; the cell lumina stellate, the trigones 
very large, with convex sides, mostly confluent ; cells of the apical 
portion 18/i in diameter, of the median portion 30/i X 2<V, and of 
the base 54/i X 25/i, a yitta not differentiated; cuticle faintly 
verruculose: underleaves imbricated, attached in a recurved line, 
subquadrate, 0.8 mm. to 1 mm. long from the line of attachment, 
the base cordate, the auricles large, undulate, rounded or toothed, 
the lateral margins sinuate, often with short teeth, the apex undu- 
late to definitely two- to four-lobed, occasionally toothed, with 
lunulate to acute sinuses, the cells as in the leaf: leaves of the 
flagelliform branches large, plane, 0.25 mm. - 0.33 mm. long X 0.24 
mm. broad at the base, ovate, the apex acute to shortly two- to 
three-toothed, the cells thick- walled as in the leaf : female branches 
occasional, solitary* one to several on a stem; the bracts and 
bracteoles similar; the outermost series short, ovate, 0.64 mm. 
long X 0.43 mm. wide, the apex entire; the intermediate series 
long-ovate, 1 mm. - 1.6 mm. long X 0.6 mm. - 0.8 mm. wide, the 
apex bifid or trifid with short teeth, the cells of the internal portion 
rectangular in outline, 45/x - 72/z X 22/t, those of the margin 
smaller, the trigones conspicuous, the walls unequally thickened; 
the innermost series similar, larger, one-fifth to one-sixth divided 
into two or three crenulate teeth, six to ten cells long, four to six 
cells broad at the base, the lateral margins crenulate, the cells 
45/A - 72/x X 22/i, trigones conspicuous, walls unequally thickened : 
perianth (immature) mouth ciliate, laciniate, the cilia and laciniae 
six to twelve cells long: male branches and sporophyte not seen. 

HABITAT : In tufts or scattered among mosses on logs and rocks 
in forests. 

The species is best characterized by its large size, and the deep 
yellow-brown to dark brown color; the long stems with densely 
imbricated, tridentate leaves with undulate to toothed auricles and 
leaf cells with stellate cell lumina and very large, rounded trigones ; 
and the large, subquadrate underleaves with strongly cordate bases, 
undulate, entire or toothed lateral margins and undulate to two- to 
four-lobed apices ; and the flagelliform branches with large leaflets. 
(FIGURES 46, nos, 1-10; 47, nos. 1-20). 

STEPHANI included this species in his Cordistipula subdivision 
(1886, 247), but later (1908, 500) transferred it to the Appendi- 
culata group. The leaves are large and often somewhat falcate. 



FIG. 46. Bazzania falcata (Lindenb.) Trevis. 1. Portion of leaf at- 
tached to stem, X 20. 2. Leaves, X 30. 3. Dorsal lobes of leaves, X 30. 
4. Apices of leaves, X 30. 5. A cell from the apical portion of a leaf, X 350. 
6. Cells from the apical tooth of a leaf, X 260. 7. Cells from the dorsal 
margin of a leaf, X 260. 8. Cells from the ventral margin of a leaf, X 260. 
9. Cells from the basal portion of a leaf, X 260. 10. Portion of a cross 
section of a stem, X 260. Nos. 1 and 2 drawn from plants collected by Bro. 
CLEMENT in Cuba; 3-10 from a portion of the type material. 



M. Fulford 132 Bazzania 

The ventral auricle or appendage (FiG. 46, nos. 1-2 and FIG. 47, no. 
1 ) was quite distinct on all of the specimens examined. Although 
it varies in size on different leaves, even on the same plant, it is 
nevertheless conspicuous. The cells are similar to those of the 
ventral edge of the leaf and the margin is undulate and usually 
set with one or more teeth. These teeth are short, broadly tri- 
angular and one to three cells high, acute or in some instances 
acuminate by the projection of a two-celled point. The leaves 
sometimes have the dorsal margins incised as indicated by 
LINDENBERG and GOTTSOHE (1851, pi. 17, fig. 4) and FiG. 46, 
no. 3; 47, no. 11. 

The leaf cells are relatively small. The trigones are very large, 
with convex sides, and appear rounded. They often become coal- 
esced or are separated only by narrow pits so that the cell lumina 
are usually stellate. In some leaves, especially on plants from very 
mesophytic situations or in the youngest leaves of the stem, the 
trigones are not quite so large and are rarely coalesced, so that 
the thin cell walls may be conspicuous. 

The underleaves are especially distinctive. They are more or 
less subquadrate, often longer than broad, deeply cordate, and at- 
tached in a recurved line. The recurved line of attachment is short 
and the base of the underleaf large and strongly cordate, so much 
so, that the basal auricles often overlap. The lateral margins are 
undulate, sometimes angled or shortly toothed, and the apex is 
rounded, undulate to two-, three-, or four-lobed or -toothed. 
STEPHANI'S drawing (I cones, Mastigobryum no. 309) of a part of 
a plant of the type, from Guadeloupe, shows an underleaf with 
undulate lateral margins and a shallow-lobed apex. There is a 
blunt tooth on one of the auricles. In another drawing, no. 415, 
also from the plants collected in Guadeloupe, the several under- 
leaves are undulate to lobed along the margins and apex. 

The flagelliform branches differ from those of the species of 
the preceding Sections in that the leaflets are much longer and 
always broader than the stem. This condition is also found in 
several other species of the Appendiculatae. 



FIG. 47. Bazzcmia falcaba (Lindenb.) Trevis. 1. Ventral view of 
stem and underleaf, X 30. 2. Underleaf, X 30. 3. Cells from the apical 
portion of an underleaf, X 260. 4. Female bract of the outermost series, 
X 30. 5. Female bract of an intermediate series, X 30. 6. Cells of an apical 
tooth of this bract. X 30. 7. Apical tooth of a bract of the innermost series, 
X 100. 8. Cells from the central portion of a bract of this series, X 350. 
9. One of the laciniae of the perianth mouth, X 100. 10. Underleaf, X 30. 
11. Basal portion of a leaf to show dorsal and ventral lobes, X 30. 12. Ventral 
appendage of a leaf, X 30. 13. A cell from the apical portion of a leaf, X 400. 
14. A leaf, X 15. 15. Ventral appendage of a leaf, X 15. 16. A cell from the 
apical portion of a leaf, X 400. 17. Underleaves, X 15. 18. Ventral auricles 
of leaves on stem, X 15. 19. A cell from the apical portion of a leaf, X 400. 
20. Underleaves, X 15. No. 1 from a plant collected by Bro. CLEMENT in 
Cuba; 2-9 from a portion of the type, from Guadeloupe; 10-13 from a portion 
of the type of M. tocutianum, from Trinidad ; 14-17 from a portion of the type 
of B. ancistrodes, from Peru; 18-20 from a portion of the type of M. armatum 
from Brazil. 



T>4" 

w 




17 



M. Fulford _134 Bazzania 

The female branches were old, poorly developed and eroded. 
The bracts of the intermediate and innermost series are character- 
ized by being one-fifth or less divided into two or three rather 
broad, crenulate teeth (see FIG. 47, nos. 4-8). The perianths 
were immature and so poorly preserved that the details of the 
mouth were the only characteristic observable. The mouth is 
divided into cilia and narrow laciniae, six to twelve cells long as 
is seen in FIG. 47, no. 9. 

LINDENBERG, in the Synopsis Hepaticarum (G. L. & N., 1844-47, 
231), records the plant from Guadeloupe and also from Nepal. 
STEPHANI (1908, 500) listed the Guadeloupe specimen and added 
that the plant from Nepal naturally does not belong here. The 
type from Guadeloupe (Hb. HOOKER) and a specimen from Nepal 
(LEHMANN) have been examined and seem to be identical, but 
more material should be studied before recording the species 
definitely from India. 

B. ancistrodes from the mountains of Peru should also be in- 
cluded under B. falcata (see FIG. 47, nos. 14-17). The plants are 
not so robust but the leaf shape, ventral auricles, and the under- 
leaves show no characteristics different from this species. Most 
of the trigones of the leaf cells are not quite so large, so that the 
thin cell wall is often conspicuous, and the brown pigmentation is 
not quite so deep. The presence of both smaller trigones and 
lighter color on one plant lead one to suspect that they are the 
results of the effect of the local environment and should not, there- 
fore, be considered of specific importance. The leaves of the 
flagelliform branches are large. 

M. armatum from Brazil (see FlG. 47, nos. 18-20) is of the 
color and size of B. ancistrodes from Peru. However, the trigones 
are larger, and are intermediate in size between those of the leaves 
of B. ancistrodes and M. falcatum. The leaves of the flagelliform 
branches are large. 

Plants collected by CRtfGER in Trinidad and described by 
STEPHANI under the manuscript name of M. tocutianum of 
GOTTSCHE are identical with this species (see FIG. 47, nos. 10-13). 
STEPHANI suggested (1886, 236) that the species was identical with 
M. falcatum of Guadeloupe and St. Vincent but he later (1908, 
500) recognized its validity. BESCHERELLE (1893 f 186) recognized 
M. tocutianum and included M. falcatum as a synonym. 

The species can readily be distinguished from B. asperistipula. 
Plants of B. falcata are strongly pigmented with brown, even in 
the younger portions, and most of the leaves are connivent, while 
plants of B. asperistipula are light yellow-green and the leaves are 
plane and subimbricated ; the appendages of the leaves of B. falcata 
are rounded to oblong and the margins are entire, undulate, and 
set with one or more short teeth, while the appendages of the 
leaves of B. asperistipula are usually large, oblong, and the margins 
are undulate, lobed, and the lobes are crenulate to shortly dentate ; 
the leaf -cell pattern of the two species is entirely different, the cell 



Bazzania 135 Appendiculatae 

lumina of the former are stellate and the trigones are large, with 
bulging sides, and confluent, or separated by narrow pits, while 
those of the latter are angular-rounded, the walls are thin and the 
trigones are small but conspicuous and very rarely are coalesced. 
The underleaves of both species are approximately the same size 
and shape and have the same mode of attachment to the stem. The 
principal difference is in the configuration of the margins. In B. 
falcata the lateral margins are entire or rarely short-toothed and 
the apex is rounded-undulate to two-, three-, or four-lobed or 
-toothed ; in B. asperistipida the whole margin is undulate to lobed, 
and crenulate to serrate, and has occasional short cilia. The 
female bracts and the mouths of the perianths of the two species 
are also different. The cell walls are thin and the laciniae long 
with ciliate or dentate margins in B. asperistipula, while in B. 
falcata the cell walls are unequally thickened, and the divisions of 
the apex are in the form of rather short, crenulate teeth. The 
laciniae of the perianth are also shorter in B. falcata. 

DISTRIBUTION : Cuba: Top of Gato Hill, 100 m,, Bro. Cl&nent 1908 
(Y, NY). Guadeloupe: without locality, Hooker Hb., the type (NY). 

Puerto Rico: Sierra de Naguabo 465-700 m., Shafer 3774, 3781 (NY). 

St. Vincent: Soufriere, Elliott 226, cited by Spruce (1895, 356); 
without locality or collector's name, Hooker Hb. (NY). Trinidad: 
Tocouche, Criiger, the type of M. tocutianum (G). Brazil : subtropics, 
Ule, the type of M. armatum (H). Peru : Mt. Campana, Spruce, Hepat. 
Sprue., the type of B. wncistrodes (H, NY). Ecuador : Cayambe, Jame- 
son (NY); Quito, Jameson (NY). 

REFERENCES: Lindenberg & Gottsche (1851, 107. pi. 17); Spruce (1895, 
356); Mitten (1861, 105); Bescherelle (1893, 186); Cook (1904, 13); Stephani 
(1908, 500; Iccnnes, Mastigobryum nos. 306, 309, 313, 372, 415). 

37. Bazzania Hookeri (Lindenb.) Trevis. Mem. 1st. Lomb. 13: 

414. 1877. 

Mastigobryum Hookeri Lindenberg, in G. L. & N., Syn. Hep. 226. 1845. 
Mastigobryum superbum Montagne, Ann. Sci. Nat. Ser. IV. Bot. 5: 349. 1856. 
Bazzania superba Trevis. loc. cit. 

Bazzania vincentina var. subrectifolia Spruce ms. p.p., Hep. Sprue. 
Bazzania flavicans Spruce, Trans. & Proc. Bot. Soc. [Edinburgh] 15: 377. 1885. 
Mastigobryum flavicans Stephani, Spec. Hep. 3: 529. 1909. 
Mastigobryum Braunianum Stephani, op. cit. p. 521. 
Mastigobryum guadaloupenae Stephani, op. cit. p. 518. 
Mastigobryum verrucosum Stephani, op. cit. p. 524. 

Mastigobryum Douini Stephani, in Herzog, Biblioth. Bot. 87: 224. Fig. 164, 
a-c. 1916. 

Plants medium size to robust, in deep tufts, olive-green to 
brown, becoming deeply pigmented in the older portions : stems to 
10 cm. long, with leaves to 4.5 mm. in width, prostrate to suberect ; 
lateral branches infrequent, diverging at a wide angle ; flagellif orm 
branches frequent, long; rhizoids colorless, present on the bases 
of the leaves of flagellif orm branches: the line of leaf insertion 
curved in its upper part, the dorsal end curved downward forming 
a hook; leaves approximate to imbricated, mostly plane, unsym- 
metrically ovate, becoming falcate, 2 mm. - 2.5 mm. long, 1 mm. - 




FIG. 48. Bazzania Hookeri (Lindenb.) Trevis. 1. Portion of a plant, 
ventral view, X 15. 2. Portion of a leaf and stem, dorsal view, X 30. 
3. Leaves, X 15. 4. A tooth of a leaf, X 310. 5. A cell from the apical portion 
of a leaf. X 400. 6. Underleaves, X 15. 7. Leaves, X 15. 8. A tooth of a 
leaf, X 310. 9. A cell from the apical portion of a leaf, X 400. 10. Under- 
leaf, X 15. Nos. 1-6 drawn from a portion of the type material; 7-10 from a 
portion of the type of if. superbum. 



Bazzania 137 Appendiculatae 

1.5 mm. broad at the base, narrowed a little to the transversely 
truncate, tridentate apex ; the dorsal margin strongly arched from 
a cordate base, extending the width of the stem and often beyond, 
the ventral margin straight to a little concave, the base dilated, 
forming an inconspicuous, entire auricle, the apex more or less 
equally three-toothed, the teeth broad, eight to ten cells long and 
broad, the sinus lunulate, the margins entire ; leaf cells thin- walled, 
the cell lumina angular-rounded, the trigones large, with convex 
sides, often coalesced, the cuticle verruculose; cells of the apical 
portion 16/A - 20/x in diameter, those of the median portion longer, 
those of the base 52/x - 64/x X 24/*, a vitta not differentiated : under- 
leaves approximate to imbricated, round-quadrate in outline, 
broader than the stem, attached in a recurved line, to 1.2 mm. long 
and broad, the base deeply cordate, the auricles large, not over- 
lapping, entire, the lateral margins and apex entire, repand or irreg- 
ularly and bluntly toothed or lobed, the cells as in the leaf : leaves 
of the flagelliform branches broader than the stem, spreading, to 
0.3 mm. long, broadly ovate, the apex acute to shortly bifid : sexual 
branches not seen. 

HABITAT : On soil and tree bases. 

The distinguishing characteristics of the species are its medium 
to large size and brownish green color; the hooked line of leaf 
insertion; the broad leaves with broad teeth and broad shallow 
sinuses, and only slightly expanded ventral auricles; thin-walled 
cells with conspicuous trigones; and the large, round-quadrate 
underleaves with entire, repand margins and strongly cordate 
bases. (FIGURES 48, nos. 1-10 ; 49, nos. 1-16) . 

The original material is made up of large plants which are 
strongly pigmented with brown. The leaves are large, spreading, 
and are broad and transversely truncate at the apex. The teeth 
are always broad, more or less equal, equilateral, and are separated 
by broad, shallow, acute sinuses, as seen in FIG. 48, nos. 1, 3 and 4. 
The cells are thin-walled and have large trigones with convex sides 
which often become coalesced (FiG. 48, nos. 4 and 5). The cuticle 
is faintly to strongly verruculose and often is extremely warty on 
the basal portions of the leaves and underleaves where the indi- 
vidual deposits sometimes tend to be oblong. The ventral appen- 
dage or auricle of the leaf, which is one of the distinguishing 
characteristics of the Section Appendiculatae, is never conspicu- 
ously developed in B. Hookeri. While many of the leaves show 
scarcely any dilation or indication of appendages, others have it 
developed to the extent that it is readily recognizable (contrast 
nos. 1, 2 and 3, FIG. 48). 

The underleaves are very large, round-quadrate, with well 
developed basal auricles which sometimes overlap. The margins 
are repand, entire, with occasional slime papillae in the depressions. 
The line of attachment is recurved. 

The presence of occasional appendages, together with the pat- 
tern of the trigones of the leaf cells, the hook-formed line of leaf 
insertion, and the large, round-quadrate, auricled underleaves 




FIG. 49. Bazzania Hookeri (Lindenb.) Trevis. 1. Portion of a plant, 
ventral view, X 15. 2. Ventral auricles of leaves on stem, X 30. 3. A cell 
from the apical portion of a leaf, X 400. 4. Underleaves, X 15. 5. Portion 
of a plant, ventral view, X 15. 6. Portion of a leaf and stem, dorsal view, 
X 30. 7. Ventral auricle of leaf on stem, X 30. 8. A cell from the apical 
portion of a leaf, X 400. 9. Underleaf, X 15. 10. Portion of a stem, ventral 
view, X 15. 11. Ventral auricle of a leaf, X 30. 12. A cell from the apical 
portion of a leaf, X 400. 13. Underleaf, X 15. 14. Portion of a plant, ventral 
view, X 15. 15. Portion of a plant, ventral view, X 15. 16. A cell from the 
apical portion of a leaf, X 400. Nos. 1-4 drawn from a portion of the type 
of M. JBraunianum, from Venezuela; 5-9 from a portion of the type of Af. 
verrueosum, from Ecuador; 10-13 from a portion of the type of M. gtutdctr 
loupense, from Guadeloupe; 14 from a portion of the type of B. vincentina 
var. subrecUfolia, from Ecuador; 15-16 from a portion of the type of B. 
flavicans, from Ecuador. 



Bazzania 139 Appendiculatae 

which are attached in a recurved line, furnish sufficient proof of 
the close relationship of this species with the other American 
members of the Section. STEPHANI included the species and 
several of its synonyms among the Section Cordistipida. 

Plants collected by UNDERWOOD, no. 1485, in Jamaica are 
smaller than the type, are light brown in color, and the under- 
leaves are longer than broad and entire along the margins. The 
other characteristics are typical of the species, including the large 
leaves of the flagelliform branches. 

Mastigpbryum superbum (FiG. 48, nos. 7-10), from Peru, and 
M. Braunianum (FiG. 49, nos. 1-4), from Venezuela, are identical 
with the type, M. Hookeri. B. flavicans (FiG. 49, nos. 15-16), 
from Ecuador, is not so deeply pigmonted and the teeth of the 
leaves are more acute and tend to be a bit longer than broad and 
more irregular, but these variations are neither constant nor of 
sufficient degree to characterize a distinct species. 

M. guadaloupense (FiG. 49, nos. 10-13), from Guadeloupe, is 
slightly more robust than M. Hookeri. Some of the teeth of the 
leaves are sharply pointed, but leaves with the shorter, broader 
teeth are not rare. The trigones are large and most of the thin 
cell walls have been obliterated by the deposition of secondary 
thickenings, so that the trigones have become coalesced. The 
ventral auricles of most of the leaves are much enlarged but they 
do not possess teeth or appendages. The cuticle is very strongly 
verruculose with large, often oblong warts. 

M. verrucosum (FiG. 49, nos. 5-9), from Ecuador, is very 
similar to M. guadaloupense in that the cuticle is very strongly 
verruculose, with oblong, wart-like protuberances, particularly in 
the older leaves and underleaves of the stem. The leaves are, for 
the most part, strongly three-toothed, but they are broad at the 
apex, as are those of the species mentioned above. M. Douini from 
Bolivia has these same characteristics. The only conspicuous 
difference between M. guadaloupense , M. verrucosum, M. Douini 
and the type specimen of M. Hookeri is this distinctly warty 
cuticle. Since these protuberances are present, but in lesser degree, 
in M. Hookeri it seems justifiable to conclude that this variation 
represents another of the expressions of a single species. 

Plants distributed by SPRUCE, from Ecuador, under the manu- 
script name B. vincentina var. subrectifolia in the Hepaticae 
Spruceana are also identical with M. Hookeri. 

The species is quite similar to B. falcata. However, in B. fal- 
cata the teeth of the leaves while broad, are not so large, the 
trigones are larger and more rounded, the ventral auricles are more 
pronounced and usually exhibit ventral appendages, and the under- 
leaves usually have teeth or appendages on the basal auricles. 

DISTRIBUTION: Cuba: Sierra Maestra, Bro. Clement 351 (NY). 
Gaudeloupe: without locality, Parker 26, the type (NY) ; without local- 
ity, 1'Herminier (NY) ; without locality, Marie, the type of M. guadaloupeme 
(H); Soufriere, Duss 320, as M. portoricense (NY). Jamaica : Blue 
Mountain Peak, Underwood 1485 (NY). Tr i ni dad : Mt. Tocouche, 



M. Fulford 140 Bazzania 

Birch (NY). Brazil: Caldas, G. A. Lindberg (NY). Bolivia: 
Yungas, Rusby 3023 (NY); Tablas, Herzog, the type of M. Douini (L). 
Ecuador: Mt. Tunguragua, Spruce, Hepat. Sprue., the type of M. ver- 
rucosum (H) ; Mt. Tunguragua, Spruce, Hepat. Sprue., the type of B. flavicans 
(H) ; the same, as B. vincentma, var. aubrectifolia p.p. (NY) ; without locality, 
Fraser, cited by Stephani (1908, 499). Guiana: Batava, Suringar, 
cited by Stephani (1909, 524). Peru : Curzco, without collector's name, 
Steph. Herb, as M. SMimicmum (H). 

REFERENCES: Lindenberg & Gottsche (1851, 77. pi. 14); Gottsche (1864, 
140) ; Stephani (1908, 499; 1909, 523; Icones, Mastigobrywm nos. 158, 312, 377, 
384, 385, 405). 

38. Bazzania robusta Spruce, Trans. & Proc. Bot. Soc. [Edin- 
burgh] 15: 378. 1885. 

Mastig obryum robustum Stephani, Spec. Hep. 3: 501. 1908. 
Mastigobryum bolivianum Stephani, in Herzog, Biblioth. Bot. 87: 223. fig. 
164, f-h. 1916. 

Plants large, in deep tufts, yellow-brown, becoming deeply 
pigmented in the older portions : stems robust, 10 cm. or more in 
length, with leaves to 5 mm. broad ; in longitudinal section the cells 
mostly 0.17 mm. long, the cortical shorter, both averaging 20/x in 
diameter, the vertical walls strongly thickened and containing fre- 
quent pits, the end walls thin; lateral branches frequent, 1 cm. or 
more apart, diverging at a wide angle; flagelliform branches fre- 
quent, long ; rhizoids colorless, present on the leaves of the flagelli- 
form branches : the line of leaf insertion curved in its upper half, 
the dorsal end recurved forming a hook ; leaves densely imbricated, 
strongly deflexed when dry, unsymmetrically ovate, falcate, 2.5 
mm. - 3.5 mm. long, 1.5 mm. broad at the base, narrowed to the 
mostly transversely truncate, tridentate apex; the dorsal margin 
arched from a cordate base, extending across the stem and beyond, 
the ventral margin strongly concave, the base conspicuously auri- 
cled, the dilation large, undulate, lobed, entire or with an occasional 
tooth; the apex deeply three-toothed, the teeth spreading, long, 
acute, to acuminate, eight to ten cells long, four to six cells broad 
at the base, the sinuses deep, lunulate to acute, the margins entire, 
undulate; the cell lumina stellate, the trigones very large, with 
concave sides, the cell walls thin, often obliterated by the enlarging 
trigones, the cuticle faintly verruculose ; cells of the apical portion 
and dorsal base averaging 24^ in diameter, those of the basal por- 
tion 32/x - 60/x long X 24/x wide, a vitta not differentiated : under- 
leaves imbricated, subquadrate-rounded, attached in a recurved 
line, broader than the stem, averaging 0.6 mm. long and broad, the 
base cordate, the auricles mostly rounded, the margins entire, un- 
dulate, lobed or faintly toothed, the lateral and apical margins 
mostly undulate, faintly lobed, obscurely toothed to entire, the cells 
as in the leaves : leaves of the flagelliform branches large, broader 
than the stem, ovate, acute or shortly bifid: female branches oc- 
casional, solitary, one to several on a stem, the bracts and bracteoles 
ovate; the outermost series short, bifid; the intermediate and 
innermost series progressively longer, one-fourth to one-sixth 
divided into two or three slender, serrate to cjliate laciniae, the 
cells to 64/x long, thin-walled, the lateral margins serrate, ciliate 




FIG. 50. Bazzania robusta Spruce. 1. Portion of a plant, ventral 
view, X 15. 2. Portion of a leaf and stem, dorsal view, X 30. 3. Leaf, X 15. 
4. Tooth of a leaf, X 310. 5. A cell from the apical portion of a leaf, X 400. 
6. Underleaves, X 15. 7. Female bract of the outermost series, X 30. 
8. Female bract of the innermost series, X 30. 9. A lacinia of this series, 
X 90. 10. Portion of the mouth of the perianth, X 90. 11. A lacinia of the 
mouth of the perianth, X 310. Nos. 1-11 drawn from a portion of the type. 



M. Fulford 142 Bazzania 

or even shortly laciniate ; perianth to 6 cm. long, the mouth ciliate 
laciniate, the cells to 64/i long, thin-walled : male branches not seen. 

HABITAT: In deep ascending tufts, on decaying logs and rocks 
and also on branches of living trees in humid forests. 

The distinguishing characteristics of the species are its large 
size and deep brown color; the falcate leaves with large, undulate, 
entire ventral auricles, large spreading teeth, and large, angular- 
rounded to stellate cell cavities with large, rounded trigones; and 
the large, subquadrate-rounded underleaves with auriculate bases, 
and undulate, faintly lobed to obscurely toothed margins. (FiG. 
50, nos. 1-11). 

The species is distinguished from B. falcata because of the 
smaller, less rounded trigones and larger cell lumina of its leaves 
and underleaves; from B. Hookeri by its larger leaf cells and 
falcate leaves with sharp teeth ; and from B. Liebmanniana by its 
larger, more acute teeth and the larger leaf cells. 

DISTRIBUTION : Brazil: Pamire* along the Rio Negro, Spruce, Hepat. 
Sprue., the type (NY); Prov. Sao Paulo, Hj. Mosen (H). Bolivia: 
Santa Anna, Williams 2161 (Y, NY) ; Corani, Herzog 5072, the type of M. 
Bolivianum (L). British Guiana: near Baitica Grove, Jenman (NY). 

REFERENCES: Stephani (1924, 466; Icones, Mastigobryum nos. 308, 311). 

39. Bazzania Liebmanniana (Lindenb. & Gottsche) Trevis. Mem. 
1st. Lomb. 13: 414. 1877. 

Mastigobryum Liebmannianum Lindenberg & Gottsche, in G. L. & N., Syn. 
Hep. 719. 1847. 

Plants medium size, light brownish green becoming darker in 
the older portions: stems slender, 5 cm. or more in length, with 
leaves to 4 mm. broad ; lateral branches diverging at a wide angle ; 
flagelliform branches frequent; rhizoids colorless, present on the 
leaves of the flagelliform branches : the line of leaf insertion curved 
in its upper part, the dorsal end recurved forming a short hook; 
leaves imbricated, strongly deflexed when dry, unsymmetrically 
ovate, ascendent, 1.5 mm. - 2.5 mm. long, 1 mm. - 1.5 mm. broad 
at the base, narrowed to the transversely to obliquely truncate, 
tridentate apex ; the dorsal margin strongly arched from a cordate 
base, extending across the stem and beyond, the ventral margin 
straight to concave, the base conspicuously auricled, the dilation 
undulate, lobed, or occasionally toothed, the apex three-toothed, the 
teeth variable, four to six cells long, four to six cells broad at the 
base, the sinus lunulate, the margins entire; the cell lumina 
angular-rounded, the cell walls thin, the trigones large, with con- 
vex sides, often becoming coalesced, the cuticle faintly verruculose ; 
cells of the apical portion and dorsal base averaging 20/x, those of 
the median portion larger, those of the basal portion 40//, - 56/x 
X 24/i, a vitta not differentiated : underleaves imbricated, subquad- 
rate, attached in a recurved line, broader than the stem, 0.6 mm. - 
0.85 mm. long and broad, the base cordate, the lateral margins 
repand, sometimes with a large tooth, the apex variously lobed or 
toothed, the cells as in the leaf : leaves of the flagelliform branches 
scale-like, to 0.2 mm, long, ovate, the apex acute to shortly bifid: 
sexual branches and sporophyte not seen. 




an' 



FIG. 51. Bazzania Liebmanniana (Lindenb. & Gottsche) Trevis. 
1. Portion of a plant, ventral view, X 15. 2. Portion of a leaf and stem, X 30.' 
3. A tooth of a leaf, X 312. 4. A cell from the apical portion of a leaf, X 400. 
5. Ventral bases of leaf on stem, X 30. 6. Underleaves, X 15. 7. Cells from 
the lateral margin of an underleaf, X 310. Nos. 1-7 drawn from a portion 
of the type. 



M. Fulf ord 144 Bazzania 

HABITAT : Not given. 

The distinguishing characteristics of the species seem to be its 
size and color; the ascendent, often falcate leaves conspicuously 
narrowed toward the apex; the short teeth, the thin- walled cells 
with conspicuous trigones and angular-rounded lumina; the sub- 
quadrate underleaves, cordate at the base, with repand margins 
which are occasionally toothed ; and the flagelliform branches with 
the typical scale-like leaves. (FiG. 51, nos. 1-7). 

I believe that this species is a variation belonging to one of the 
other species included in this Section, but the portion of the type 
in the STEPHANI Collections in the Farlow Herbarium is so frag- 
mentary that it has been impossible to get an adequate concept of 
the range of variability of the LINDENBERG and GOTTSCHE species. 

It can be distinguished from the preceding species by its 
spreading leaves which tend to be narrowed in the apical region, 
the small, undulate ventral auricles with entire margins, and by 
the thin cell walls and conspicuous trigones. 

DISTRIBUTION: Mexico : Oaxaca, Liebmann, the type (H); Vera Cruz, 
Purpus 5548 (Y). Guatemala: Coban, Standley 69119 (W). 

REFERENCES: Gottsche (1863, 133, 136); Lindenberg & Gottsche (1851, 
74, 81); Stephani (1909, 523; I cones, Mastigobryum no. 392); Herzog (1938, 
19). 

40. Bazzania teretiuscula (Lindenb. & Gottsche) Trevis. Mem. 1st. 

Lomb. 13: 414. 1877. 

Mastigobryum teretiusculum Lindenberg & Gottsche, in G. L. & N., Syn. Hep. 

720. 1847. 
Bazzania keteroclada Spruce, Trans. & Proc. Bot. Soc. [Edinburgh] 15: 379. 

1885. 

Mastigobryum heterocladum Stephani, Spec. Hep. 3: 502. 1908. 
Bazzania spinigera Spruce, op. cit. p. 380. 
Mastigobryum spinigerum Stephani, op. cit. p. 500. 
Bazzania humifusa Spruce, op. cit. p. 379. 
Mastigobryum humifusum Stephani, Hedwigia 44: 225. 1905. 
Bazzania conchopkylla Herzog, Rev. Bryol. et Liche*n. 11: 20. 1938. 
Mastigobryum incisostipulum Stephani, in Herzog, Biblioth. Bot. 87: 225. 

Fig. 165, c-d. 1916. 

Plants medium size, in tufts or mats, greenish brown, deeply 
pigmented with brown in the older portions: stems to 8 cm. long, 
with leaves to 3.2 mm. broad, suberect; stem cells in longitudinal 
section elongate, averaging 0.17 mm. long, the cortical shorter, 
both averaging 20/u, in diameter, the vertical walls uniformly 
thickened, containing frequent pits, the end walls thin: lateral 
branches 5 mm. or more apart, diverging at a wide angle ; flagelli- 
form branches frequent; rhizoids colorless, few, on the bases of 
the leaves of the flagelliform branches: the line of leaf insertion 
curved in the upper half, the dorsal end recurved forming a hook ; 
leaves approximate to imbricated, deflexed, strongly so when dry, 
unsymmetrically ovate, straight to subfalcate, 1.2 mm. - 2.2 mm. 
long, to 1 mm. broad at the base, narrowed to the more or less 
obliquely truncate, tridentate apex; the dorsal margin strongly 
arched from a cordate base, extending the width of the stem and 
beyond, the ventral margin concave, the base dilated, the auricle 




FIG. 52. Bazzania, teretiiiseula (Lindenberg & Gottsche) Trevis. 

I. Portion of a plant, ventral view, X 15. 2. Portion of leaves and stem, 
dorsal, view, X 30. 3. Leaves, X 15. 4. Cells from the apical portions of 
leaves, X 400. 5. Underleaves, X 15. 6. Portion of stem, showing auricles 
of leaves, X 15. 7. Leaf, X 15. 8. A cell from the apical portion of a leaf, 
X 400. 9. Underleaves, X 15. 10. Portion of a stem, ventral view, X 15. 

II. Underleaf, X 15. 12. A cell from the apical portion of a leaf, X 400. 
13. Underleaves, X 15. Nos. 1-5 drawn from a portion of the type, from 
Mexico; 6-9 from a portion of the type of B. humifusa,, from Peru; 10-11 from 
a portion of the type of B. otites, from Peru; 12-13 from a portion of the 
type of B. conchophylla, from Costa Rica. 



M. Fulford _146 Bazzania 

undulate, entire, appendiculate, toothed, or ciliate, the apex irregu- 
larly three-toothed, the teeth mostly spreading, acute, five to eight 
cells long, four to six cells broad at the base, the sinuses deep, 
lunulate, the margins entire, repand ; leaf cells thin-walled, the cell 
lumina angular-rounded, the trigones conspicuous, with convex 
sides, often confluent, the cuticle faintly verruculose; cells of the 
apical region and dorsal base averaging 20ji X 20/x, those of the 
interior larger, those of the base to 36/x X 20/*, a vitta not differ- 
entiated; underleaves approximate to imbricated, subquadrate in 
outline, averaging 0.7 mm. long and broad, attached in a recurved 
line, broader than the stem, the base cordate, the auricles not over- 
lapping, rounded, the margins entire, or dentate with one or two 
broad teeth, or ciliate by a row of two or three cells, the lateral 
margins convex, entire or rarely with one or two teeth, the apex 
deeply four-lobed or -toothed, the sinuses narrow, acute, the cells 
and trigones as in the leaf: leaves of the flagelliform branches 
scale-like, ovate, acute to shortly bifid : female branches occasional, 
solitary, one to several on a stem, the bracts and bracteoles similar, 
ovate (immature), the outermost series small, shortly bifid, the 
intermediate series larger, to one-sixth divided into three laciniae, 
the cells mostly 20/x X 16/x, with thick walls, the lateral margins 
serrate to short ciliate; the apical portion of the innermost series 
divided into three, serrate, ciliate laciniae, the cells mostly 
36 ^ X 16/i, the cell walls thin : male branches and perianth not seen. 

HABITAT: On rocks and bases of trees on shaded slopes in 
moist forests. 

The distinguishing characteristics of the species are its medium 
size and greenish brown color; the subfalcate, tridentate leaves 
with thin cell walls and distinct trigones; the enlarged ventral 
auricles with wavy, entire, toothed or appendiculate margins ; and 
the subquadrate, cordate, underleaves with definitely four-lobed or 
-toothed apices, and approximate, rounded, basal auricles, having 
entire or occasionally toothed or ciliate margins. (FIGURES 52, 
nos. 1-13; 53, nos. 1-20). 

The plants show much variation both as to size and to con- 
figuration of the parts. On some areas of a stem the leaves and 
underleaves may be densely imbricated while on others they may 
be approximate or even distant. Many of the leaves tend to be 
narrow-elongate in the upper part, but shorter leaves are not 

FIG. 53. Bazzania teretiuscula (Lindenb. & Gottsche) Trevis. 1. Por- 
tion of plant, ventral view, X 15. 2. Portion of a leaf and stem, dorsal view, 
X 30. 3. Tooth of a leaf, X 310. 4. A cell from the apical portion of a leaf, 
X 400. 5. Cells from the basal portion of a leaf, X 312. 6. Outline of basal 
auricles of leaves, X 30. 7. Underleaves, X 15. 8. Female bract of an outer 
series, X 30. 9. Female bract of the intermediate series, X 30. 10. A lacinia 
of a bract of the innermost series, X 100. 11. The tip of the same lacinia, 
X 310. 12. Portion of a plant, ventral view, X 15. 13. A cell from the apical 
portion of a leaf, X 400. 14. Auricles of ventral bases of leaves, X 30. 
15. Underleaves, X 15. 16. Female bract of outer series, X 30. 17. Female 
bract of intermediate series, X 30. 18. A lacinia of a bract of this series, 
X 310. 19. A lacinia of a bract of the innermost series, X 100. 20. Cells of 
this same lacinia, X 310. Nos. 1-11 drawn from plants of the type of B. 
heteroclada, from Peru; 12-20 from plants of the type of B. spinigera, from 
Brazil. 



M. Fulford 148 Bazzania 

uncommon. The cell walls are thin, and in some leaves they are 
very conspicuous together with the small trigones, but in others 
(different plants of the same tuft), the trigones are much larger 
and many of them have become coalesced. There is so much 
variation in the ventral auricles of the leaves on any stem that no 
typical form can be distinguished. The ventral base is always 
dilated, but the dilation may be large or small, straight or undulate, 
with an entire margin or with one or two teeth, or several cilia 
of two or three cells. 

The underleaves also show a high degree of variation. They 
are always four-lobed or -toothed on the apical margins and are 
more or less auricled at the bases. The underleaves of well 
developed plants have auricles which are large but do not overlap. 
Some sort of tooth or appendage is usually developed near the base. 
On stems where the leaves and underleaves are only approximate 
or are distant, the basal auricles of the underleaves are smaller and 
are rounded-entire and show no indication of projections on the 
margins. They are usually four-lobed but may be only four-toothed. 

The plants of the type material, from Mexico (FIG. 52, nos. 
1-5), are robust, well developed, and show a wide range of variation 
both in the leaves and underleaves. The plants of B. heteroclada, 
collected by SPRUCE in Peru (FIG. 53, nos. 1-11), are very similar. 
Some of the teeth of the leaves are narrower and more acute, and 
there is more variation in the margins of the ventral auricles of 
the leaf (see FIG. 53, nos. 1 and 6), but these two characteristics 
are not constant, even on adjacent leaves. 

SPRUCE has also described plants with similar characteristics 
from Brazil as B. spinigera (see FIG. 53, nos. 12-20). The shape 
of the leaves is similar to that of the well developed forms of M. 
teretiusculum; the leaf cells are of approximately the same size 
and the cell walls are thin, but the trigones often tend to be some- 
what larger in most plants. Here too, the ventral auricles exhibit 
a wide variety of forms on a single stem, but those with one or two 
cilia, two or three cells long, are more abundant, although auricles 
with blunt teeth or with rounded-entire margins are not uncommon. 
The underleaves also show much variation in the cordate bases. 
One or two teeth or cilia are usually well developed and occur on 
most of the auricles, although auricles with entire margins are 
frequent. The apical margins more often are divided into four 
teeth rather than four lobes as is the more usual condition in B. 
heteroclada. 

Female branches were present in both B. heteroclada and /?. 



FIG. 54. Bazzania, acanthostipa Spruce. 1. Portion of a plant, ventral 
view, X 15. 2. Portion of a leaf and stem, dorsal view, X 30. 3. Ventral 
auricle of leaf on stem. X 30. 4. Tooth of a leaf, X 312. 5. A cell from the 
apical portion of a leaf, X 400. 6. Underleaf, X 30. 7. Female bract of the 
innermost series, X 30. 8. A part of the perianth mouth. X 90. 9. A portion 
of a lacinia of the perianth mouth, X 312. 10. Portion 01 a transverse section 
of a stem, X 312. Nos. 1-10 drawn from a portion of the type. 



M. Fulford 150 Bazzania 

apinigera. The bracts were immature and the archegonia unfer- 
tilized. While a comparison of the two (FiG. 53, nos. 8-11 and 
16-20), shows that there are minor variations, in general the two 
are very similar as to form, margin, type of laciniae and shape 
of the cells. 

SPRUCE collected plants with similar characteristics on Mt. 
Campana, Peru, among Sphagnum, and distributed them as B. 
humifusa (see FIG. 52, nos. 6-9), in the Hepaticae Spruceana. 
Although the leaves exhibit a wide variation in outline, with the 
ventral auricles less well developed, and with only occasional short 
teeth on the basal auricles of the underleaves, the plants possess no 
distinctive characteristics not present in the forms described above. 
The four lobes of the underleaves are always well developed. The 
cell walls are thin but the trigones are conspicuous. All of the 
features of the plants indicate that they grew under very moist 
or wet conditions of habitat. 

In the Herbier Boissier at Geneva there is a collection made by 
SPRUCE in 1885 on Mt. Campana, Peru, filed under M. teretiusculum, 
but on the packet in SPRUCE'S handwriting is the name B. otites 
R. S. n. sp. The plants are large, green and well developed but 
sterile, and exhibit on one part or another, all of the characteristics 
of M. teretiusculum (see FIG. 52, nos. 10-11). Most of the under- 
leaves are definitely four-lobed and some of them have in addition, 
a large lateral tooth. The basal auricles are well developed and 
occasionally appendages in the form of cilia occur. 

B. teretiuscula of SPRUCE (1885, 375), is B. longistipula. 

The plants collected by STANDLEY in Costa Rica and distributed 
as B. conchophylla Herzog, also belong to this species (see FIG. 
52, nos. 12-13). They are lighter in color than many of the South 
American plants, but the leaves and underleaves have all of the 
characteristics of M. teretiusculum. 

M. incisostipulum Stephani collected by HERZOG in Bolivia also 
belongs here. The plants are robust, greenish or yellow-brown, and 
the underleaves are typically conspicuously four-lobed. 

The species can be distinguished from those preceding by the 
deeply four-lobed or -toothed apical portions of the underleaves and 
the well developed ventral auricles of the leaves. 

DISTRIBUTION : Mexico: Vera Cruz, Hacienda de Mirador, Liebmann, 
the type (G). Costa Rica: La Estrella, Standley 39426 (W); San 
JOSH, Standley 41629 (W) ; Cartago, Standley 50, 884 (W). Brazil: Rio 
Taruma at the Rio Negro, Spruce, Hepat. Sprue., the type of B. spinigera 
(NY). Bolivia: Comarapa, Herzog, the type of M. incisostipulum (L). 
Colombia: Bogota, Apollinaire, cited by Stephani (1909, 520). 
Ecuador : Quito, Cuming, cited by Stephani (1909, 520). Peru : Mt. 
Guayrapurina, Spruce, Hepat. Sprue., the type of B. heteroclada (NY) ; Mt. 
Campana, Spruce, Hepat. Sprue., the type of B. humifusa (NY) ; Mt. Cam- 
pana, Spruce, under M. teretiusculum and B. otites Sp. (G) ; Rio Huallaga, 
Ule, cited by Stephani (1904,225). 

REFERENCES: Lindenberg & Gottsche (1851, 85. pi. 20); Gottsche (1863, 
132) ; Stephani (1909, 522, 520; Icones, Mastigobryum nos. 166, 310, 387, 402) : 
Herzog (1938, 11). 



Bazzania 151 Appendiculatae 

41. Bazzania acanthostipa Spruce, Trans. & Proc. Bot. Soc. [Edin- 
burgh] 15: 381. 1885. 

Mastigobryum acanthostipum Stephani, Spec. Hep. 3: 499. 1908. 

Plants large, in deep tufts, light brown, greenish in the younger 
portions: stems robust, to 14 cm. in length, with leaves to 5 mm. 
broad; in longitudinal section the cells mostly 0.17 mm. long, the 
cortical shorter, both averaging 18/x in diameter, the vertical walls 
strongly thickened and containing frequent pits, the end walls thin, 
walls of the cortical layer deeply pigmented; lateral branches oc- 
casional, diverging at a wide angle ; flagellif orm branches frequent, 
long: rhizoids colorless, present on the leaves of the flagellif orm 
branches: the line of leaf insertion curved in its upper half, the 
dorsal end recurved forming a hook; leaves densely imbricated, 
deflexed when dry, unsymmetrically ovate, falcate, 2.5 mm. - 3.5 
mm. long, 2 mm. broad at the base, narrowed to the mostly trans- 
versely truncate, more or less equally tridentate apex; the dorsal 
margin arched from a cordate base, extending across the stem and 
beyond, the ventral margin strongly concave, the base conspicu- 
ously auricled, the dilation very large, undulate, lobed, toothed, 
often appendiculate; the apex deeply three-toothed, the teeth 
spreading, long, broadly triangular, acute, eight to fifteen cells 
long, four to six cells broad at the base, the sinuses deep, lunulate, 
the margins entire, repand; the cell lumina stellate, the trigones 
very large, with convex sides, the cell walls thin but obscured by 
the enlarged trigones, the cuticle faintly verruculose; cells of the 
apical portion and dorsal base averaging 20/x X 20/x, those of the 
basal portion 48/x - 64/x X 24/x, a vitta not differentiated: under- 
leaves imbricated, subquadrate, attached in a recurved line, 
broader than the stem, averaging 1.2 mm. long and broad, the base 
cordate, auriculate, the auricles very large, overlapping, undulate, 
serrate, lobed, toothed, or appendiculate, the lateral margins 
coarsely toothed or lobed, and with one or more broad to narrow, 
apiculate, widely spreading teeth, the apical margin convex, undu- 
late, sometimes four-lobed, the cells and trigones as in the leaf; 
leaves of the flagelliform branches plane, ovate, to 0.22 mm. long, 
acute to shortly bifid : female branches occasional, solitary, one to 
several on a stem; the bracts and bracteoles similar, ovate; the 
outermost series short, bifid ; the intermediate series larger, to one- 
third divided into usually three, ciliate laciniae, the lateral margins 
ciliate; the innermost series to 2.2 mm. long, one-fourth divided 
into three ciliate and serrate laciniae, the cells to 64ju long, with 
thickened walls, the lateral margins serrate to long-ciliate : perianth 
to 6 mm. long, the mouth ciliate laciniate, the cells averaging 32/x 
long, thick- walled : male branches not seen. 

HABITAT : In deep ascending tufts, on rocks in moist places in 
the mountains. 

The distinguishing characteristics of the species are its large 
size and light brown color ; the falcate leaves, with large, spreading 
teeth, stellate cell cavities, with very large, nearly round trigones, 
and very large, undulate, lobed, often appendiculate, ventral 
auricles; and the very large, subquadrate, cordate underleaves, 



M. Fulford 152 Bazzania 

with large basal auricles similar to those of the leaves, and one or 
more large, widely spreading, usually long apiculate, lateral teeth. 
(FlG. 54, nos. 1-10). 

The species is distinct from all other members of the Appendi- 
culatae because of its size, the very large teeth of the leaves, and 
the long, pointed teeth of the lateral margins of the underleaves. 

DISTRIBUTION: Colombia: without locality, A. Wallace (NY). 
Bolivia : Mapiri, Rusby 3027 p.p. (NY). Peru : Mt. Campana, Spruce, 
Hepat. Sprue., the type (NY). 

REFERENCES: Stephani (Icones, Mastigobryum no. 305). 

42. Bazzania canelensis (Steph.) comb.nov. 

Mastigobryum canelense Stephani, Spec. Hep. 3: 518. 1909. 
Bazzania vincentina var. submutica Spruce, Trans. & Proc. Bot. Soc. [Edin- 
burgh] 15: 378. 1885. 
Bazzania vincentina var. subedentata Spruce ms., Hepat. Sprue. 

Plants large, in deep tufts, olive-green to yellow-brown, becom- 
ing more deeply pigmented in the older portions: stems large, to 
10 cm. or more in length, with leaves to 5 mm. broad, ascending 
to erect; stem cells in longitudinal section to 0.25 mm. long, the 
cortical shorter, both averaging 18/x in diameter, the vertical walls 
uniformly thickened and containing frequent pits, the end walls 
thin ; lateral branches rare, diverging at a wide angle ; flagellif orm 
branches frequent, short; rhizoids very abundant, on the leaves of 
the flagellif orm branches: the line of leaf insertion curved in its 
upper part, the dorsal end recurved forming a prominent hook; 
leaves approximate to imbricated, plane, becoming a little deflexed 
when dry, unsymmetrically ovate to oblong, straight, 2.5 mm. - 3 
mm. long, 1.5 mm. broad at the base, narrowed a little to the 
broadly rounded to faintly tridentate apex; the dorsal margin 
strongly arched from a cordate base, extending across the stem and 
somewhat beyond, the ventral margin straight, the base dilated, 
the auricle large, undulate, sometimes lobed, the margin entire to 
serrulate, the apex broad, rounded or undulate to three-lobed or 
-toothed, the lobes or teeth broad, short, the sinuses broad, 
lunulate, the margins entire ; leaf cells thin- walled, the cell lumina 
angular-rounded, the trigones conspicuous, sometimes becoming 
coalesced, the cuticle smooth to faintly verruculose; cells of the 
apical region and dorsal lobe averaging 16//, - 20/i, those of the 
interior larger, those of the base to 60/t X 20^, a vitta not differ- 
entiated; underleaves approximate to imbricated, subquadrate in 
outline, attached in a recurved line, broader than the stem, 
1.5 mm. - 2 mm. long, 1.5 mm. broad, the base strongly cordate, the 
auricles large, overlapping or nearly so, the margins entire, undu- 
late, the lateral margins convex, the apical margins rounded, un- 
dulate, slime papillae present in the depressions, the cells as in the 
apical portion of the leaves: leaves of the flagellif orm branches 
scale-like, ovate, acute to shortly bifid: sexual branches and peri- 
anths not seen. 

HABITAT : On tree trunks and branches, forests. 

The distinguishing characteristics of the species are its large 




FIG. 55. Bazzania canelensis (Steph.) Fulford. 1. Portion of a plant, 
ventral view, X 15. 2. Portion of a leaf and stem, dorsal view, X 15. 3. Leaf 
apices, X 15. 4. Tooth of a leaf, X 310. 5. A cell from the apical portion of 
a leaf, X 400. 6. Cells from the basal portion of a leaf, X 310. 7. Ventral 
base of a leaf with appendage, X 30. 8. Underleaf, X 15. 9. Portion of the 
apical margin of an underleaf, X 310. 10. Portion of a cross section of a 
stem, X 310. Nos. 1-10 drawn from plants of the type of B. subedentata 
Spruce, from Canelos, Ecuador. 



M. Fulford 154 Bazzania 

size and pale brownish green color; the large leaves with rounded- 
entire to faintly three-lobed or -toothed apices, large, undulate, 
entire to crenulate ventral auricles, and thin-walled cells with 
angular-rounded cell lumina and conspicuous trigones; and the 
large, subquadrate, auriculate underleaves with entire, repand to 
undulate margins. (FiG. 55, nos. 1-10). 

While the type plants of B. vincentina van submutica Spruce, 
collected by SPRUCE at Canelos have not been available for study, 
his description of the variety agrees in detail with the plants which 
he distributed under the manuscript name B. vincentina var. 
subedentata in the Hepaticae Spruceana. In addition, STEPHANI 
described the species M. canelense from material collected by 
SPRUCE from the same locality and lists B. vincentina var. submutica 
as a synonym. The plants of M. canelense in the STEPHANI Col- 
lection at Harvard are identical with those distributed by SPRUCE 
as B. vincentina var. subedentata. It would seem that the two 
varieties and the species named above are all based on plants from 
the same collection. 

The species is distinct from all other members of the Appendicu- 
latae because of its large size, the entire or obscurely three-toothed 
leaf apices, and the large underleaves with more or less entire 
margins. 

DISTRIBUTION : Ecuador: Canelos, Spruce, Hepat. Sprue., the type of 
B. vincentina var. subedentata (NY) ; the same, the type of M. canelense (H) ; 
the same, the type of B. vincentina var. submutica, cited by Spruce. 

REFERENCES: Stephani (/cones, Mastigobryum no. 379). 



Section 5. Vittatae 

The species of this Section have three-toothed leaves in which 
a vitta is conspicuously developed. The yitta is made up of several 
rows of cells which are elongate in outline, thin-walled, and with 
conspicuous trigones, and it extends from the base of the leaf to 
beyond the middle. 

Key to the Species 

1. Plants with underleaves chlorophyllose throughout; the margins variously 

lobed or toothed 43. B. Spruceana (p. 155) 

1. Plants with the underleaves hyaline at least in the upper part; more or 
less regularly four-toothed, the lateral margins entire. 
2. Cuticle of the leaves and underleaves abundantly minutely punctuate; 
underleaves hyaline throughout . . . . 44. B. Tayloriaiut (p. 157) 
2. Cuticle of the leaves and underleaves smooth to verruculose; under- 
leaves chlorophyllose in the lower part. 
3. Underleaves to one-half divided into four, equal, narrow teeth or 

lobes; medium size 45. B. convexa (p. 159) 

3. Underleaves very large, often four-toothed or -lobed in the upper 
part 46. B. Step haul (p. 162) 

43. Bazzania Spruceana Stephani, Hedwigia 32 : 213. 1893. 

Maatigobryum Spruceanum Stephani, Spec. Hep. 3: 469. 1908. 

Plants small, brownish green becoming deeply pigmented with 
brown in the older portions: stems slender, 3 cm. or more long, 
with leaves to 2 mm. broad: lateral branches frequent, 2 mm. or 
more apart, diverging at a wide angle; flagelliform branches oc- 
casional; rhizoids colorless, present on the bases of some of the 
underleaves: the line of leaf insertion curved in the upper part; 
the leaves vittate, imbricated, plane, unsymmetrically ovate, as- 
cendent, 0.7 mm. - 0.9 mm. long, 0.4 mm. broad at the base, 
narrowed a little to the mostly transversely truncate, tridentate 
apex; the dorsal margin convex from a straight base, covering 
one-half the stem, the ventral margin straight, the base scarcely 
dilated, the apex mostly equally three-toothed, the teeth small, 
acute, spreading, two to five cells long, two to six cells broad at the 
base; the sinuses shallow, lunulate, the margins entire: the leaf 
cells round-quadrate in outline, the walls uniformly thickened, the 
cell lumina rounded, trigones inconspicuous or absent, the cuticle 
very strongly verruculose ; the cells of the apical region 16/i - 22/x 
in diameter, those of the median portion larger, and those of the 
base to 48/i X 24/i, forming a distinct vitta of three or four rows of 
cells extending beyond the middle of the leaf: underleaves distant 
to imbricated, subquadrate in outline, attached in a straight line, 
a little broader than the stem, mostly 0.35/i long and broad, the 
lateral margins slightly convex, entire or occasionally with a tooth, 
the apex entire, crenate, undulate or two- to four-lobed, the cells 




FIG. 56. Bazzania Spruceana Stephani. 1. Portion of a plant, ventral 
view, X 15. 2. Portion of a leaf and stem, dorsal view, X 30. 3. A leaf, 
X 30. 4. A cell from the apical portion of a leaf, X 400. 5. Teeth from a 
leaf, X 310. 6. Cells of the vitta near the base of a leaf, X 310. 7. Under- 
leaves. 8. Cells from the apical portion of an underleaf, X 400. Nos. 1-8 
drawn from a portion of the type material. 



Bazzania 157 Vittatae 

as in the leaf: leaves of the flagelliform branches scale-like, ovate: 
sexual branches not seen. 

HABITAT : Not given. 

The distinguishing characteristics of the species are its small 
size; the plane, ascendent leaves with three small, sharp teeth at 
the apex, cells with equally thickened walls, and a distinct vitta; 
and the small, subquadrate underleaves with variously lobed 
margins. The cuticle is coarsely verruculose. (FiG. 56, nos. 1-8). 

DISTRIBUTION: Peru : Mt. Guayrapurina, Spruce, the type (H). 

REFERENCE: Stephani (Icanes, Mastigobryum no. 177). 

44. Bazzania Tayloriana (Mitten) comb. nov. 

Mastigobryum Tayloriamim Mitten, in Hooker, Bot. Ant. Voy. 2 1 . Fl. N. Zel. 
147. pi. C, fig. 5. 1854. 

Plants small, growing in depressed mats, pale, glaucous green, 
becoming deeply pigmented with brown in the older portions: 
stems slender, delicate, to 3 cm. or more in length, with leaves 
mostly 1.5 mm. broad, prostrate; stems cells in longitudinal section 
elongate, the vertical walls uniformly thickened and containing 
frequent pits, the end walls thin; lateral branches occasional, 
diverging at a wide angle, the ventral branches frequent, usually 
leafy; rhizoids not seen: the line of leaf insertion little curved in 
its upper part; the leaves approximate to subimbricated, vittate, 
spreading, plane, becoming somewhat deflexed beyond the middle, 
unsymmetrically oblong-ovate, 0.7 mm. - 1.0 mm. long, mostly 
0.5 mm. broad at the base, little narrowed to the transversely 
truncate, conspicuously tridentate apex ; the dorsal margin convex 
from a straight base which covers less than one-third the stem, 
the ventral margin nearly straight, the base scarcely dilated, the 
apex tridentate, the teeth large, spreading, eight to ten cells long, 
four to six cells broad at the base, the sinuses deep, acute to 
rounded, the margins entire; the leaf cells quadrate in outline, 
uniformly thick-walled, the cell lumina rounded, a vitta of three 
or four rows of enlarged cells with conspicuous trigones clearly 
differentiated, the cuticle very abundantly minutely punctate ; cells 
of the apical portion and dorsal base mostly 16/x X 16/x, of the 
vitta, to 32^ X 24/x, the walls thin, trigones conspicuous: under- 
leaves distant, hyaline throughout, attached in a straight line, 
subrectangular in outline, broader than the stem above, 0.22 mm. - 
0.35 mm. long, to 0.28 mm. broad above, the lateral margins nearly 
straight from a straight base, one-fourth to one-half divided into 
usually four equal, blunt teeth four to ten cells long, two to four 
cells broad, the sinuses acute, the margins entire ; the cells quadrate 
to rectangular in outline, hyaline, the walls uniform, the cuticle 
abundantly minutely punctate as in the leaves: sexual branches 
not seen. 

HABITAT : On soil among other bryophytes. 

The distinguishing characteristics of the species are its small 
size and glaucous to brownish green color; the strongly three- 
toothed leaves with small, quadrate leaf cells, and a well marked 




FIG. 57. -T- Bazzania Tayloriana (Mitten) Fulford. 1. Portion of plant, 
ventral view, X 30. 2. A tooth of a leaf, X 310. 3. A cell from the apical 
portion of a leaf, X 400. 4. Underleaves, X 30. 5. A tooth of an underleaf, 
X 310. The drawings are from the type material. 



Bazzania 159 Vittatae 

vitta; the hyaline, two- or four-parted elongate underleaves; and 
a very abundantly and minutely punctate cuticle. (FiG. 57, 
nos. 1-5). 

B. Tayloraniana can readily be separated from all of the other 
South American species because of the abundantly minutely 
punctate cuticle of the leaves and underleaves. 

DISTRIBUTION: Colombia: Bogotd, 4000', Weir (NY). New 
Zealand: North Island, Zotov (NY); North Island, Colenso, the type 
(NY) ; also Colenso 223, 1218 (NY) ; North Island, Sinclair (NY) ; Kaipara 
?, Mossman 760d (NY); without locality, Hooker, Knight, Kirk, cited by 
Stephani. Samoa: without locality, Rechinger, cited by Stephani. 

REFERENCES: Hooker (1867, 524); Stephani (1909, 533; /cones, Masti- 
gobryum no. 465). 

45. Bazzania convexa (Thunb.) Trevis. Mem. 1st. Lomb. 13: 414. 

1877. 

Jungermannia convexa Thunberg, Prod. PL Cap. 173. 1794. 

Jungermannia nitida Weber, Prodromus 43. 1815. 

Mastigobryum convexum Lindenberg, in G. L. & N., Syn. Hep. 215. 1845. 

Mastigobryum Richardianum Mitten, in Hooker, Bot. Ant. Voy. 2 s . FL N. Zel. 

147. 1854. 
Mastigobryum heterostipum Stephani, Spec. Hep. 3: 532. 1909. 

Plants scattered, or growing in depressed mats, pale, yellow- 
green : stems very slender, to 3 cm. or more in length, with leaves 
to 1.5 mm. broad, prostrate ; lateral branches few, diverging at a 
wide angle, ventral branches frequent, usually leafy, sometimes 
flagelliform: rhizoids frequent on the bases of the leaves of the 
flagellif orm branches : the line of leaf insertion little curved in its 
upper part; the leaves vittate, spreading, approximate to subim- 
bricated, nearly straight, the teeth and a little of the apical portion 
deflexed, unsymmetrically oblong-ovate, 0.7 mm. - 0.9 mm. long, 
mostly 0.35 mm. broad at the base, narrowed a little to the trans- 
versely truncate, mostly obscurely tridentate, decurved apex; the 
dorsal margin convex from a straight base which covers less than 
one-half the stem, the ventral margin nearly straight, the base 
scarcely dilated, the apex obscurely tridentate, the teeth triangular, 
one or two cells long and wide, the sinuses lunulate, the margins 
entire; the leaf cells quadrate to rectangular in outline with uni- 
formly thickened walls, the cell lumina rounded, the vitta of three 
or four rows of enlarged cells clearly differentiated; the cuticle 
verruculose; cells of the apical portion, margins and dorsal base 
averaging 16/A X 16/x, those of the vitta to 32/* X 24/i, the walls 
thin, the trigones conspicuous : underleaves distant, hyaline above, 
attached in a straight line, subquadrate in outline, averaging 0.28 
mm. X 0.28 mm., mostly one-third divided into four equal, acute 
teeth, the teeth mostly five cells long, to four cells broad at the 
base, the sinuses acute, the margins entire ; cells of two sorts, the 
chlorophyllose cells restricted to a small, interior, basal area, 
similar to those of the margins of the leaf, the hyaline cells larger, 
more or less rectangular in outline, averaging 18/x - 22/x long and 
wide, the walls uniformly thickened: leaves of the flagelliform 
branches, ovate, scale-like: sexual branches not seen. 



M. Fulford 160 Bazzania 

HABITAT: On soil, among rocks, mixed with other bryophytes. 

The distinguishing characteristics of the species are its small 
size and yellowish green color; the straight, obscurely tridentate 
leaves with slightly decurved tips and distinct vittas; and the 
equally four-toothed underleaves which are hyaline except for 
a small area at the base. (FiG. 58, nos. 1-17). 

The plants are always small and on well developed stems the 
leaf apices are shortly three-toothed (FIG. 58, no. 6), with the 
teeth and a portion of the leaf decurved. This characteristic is 
not always present however, for in many examples the leaves may 
be entirely plane and rounded-entire, or very slightly three-lobed. 
Some leaves of the more typical form can usually be found on any 
stem more than 1 cm. in length. 

Very often stems with normally developed leaves suddenly be- 
come flagelliform for a distance and then produce ordinary leaves 
again. The ventral branches are usually flagelliform but leafy 
branches are not uncommon. 

The underleaves are extremely variable. Usually they are one- 
third to one-half divided into four acute teeth but very often the 
teeth do not develop in the ordinary way or the cells of the upper 
part soon disappear, for many of the underleaves are only bluntly 
four-lobed. The cells of the chlorophyllose area, in the basal part 
of the underleaf, are similar to the cells of the apical portion of 
the leaf. The hyaline cells are usually rectangular in outline, more 
or less regularly arranged in rows, and have uniformly thickened 
walls (see FIG. 58, nos. 4, 5 and 17). This is particularly true 
of the plants of South Africa and Australia. However, one often 
finds, growing in the same patches with the plants bearing under- 
leaves of the sort described, similar plants in which the cells of 
the underleaves are more irregularly arranged, are slightly larger, 
and for the most part, have thin walls, although some of the walls 
may be thick (see FIG. 58, no. 14). Most of the plants of the type 
of M. Richardianum have this latter sort of underleaves. FiG. 58, 
no. 5 shows a tooth with large cells but with thick walls. Plants 
with underleaves which have the larger, irregularly arranged, thin- 
walled cells can likewise be found in the African material. Since 
this combination of characters is not entirely constant in any mat 
of plants they do not seem of sufficient significance to serve as a 

FIG. 58. Bazzania convexa (Thunb.) Trevis. 1. Portion of plant, 
ventral view, X 30. 2. A cell from the apical portion of a leaf, X 400. 3. An 
underleaf, showing the hyaline part, X 30. 4-5. A tooth of an underleaf, 
X 310. 6. Portion of a plant, ventral view, X 30. 7. Portion of a stem and 
leaf, dorsal view, X 30. 8. Apices of leaves, X 30. 9. A leaf, X 30. 10. A 
cell from the apical portion of a leaf, X 400. 11. An apical tooth, X 310. 
12. Cells of the vitta, X 310. 13. Underleaf, showing the hyaline portion, 
X 30. 14. Two teeth of an underleaf, X 310. 15. Portion of a plant, ventral 
view, X 30. 16. Underleaf showing the hyaline part, X 30. 17. A tooth of 
an underleaf, X 310. Nos. 1-4 drawn from the type of J. convexa from the 
Cape of Good Hope; 5 from J. convexa of Lindenberg from Peru; 6-14 from 
the type of M. Richardianum from the Straits of Magellan; 15-17 from the 
type (?) of Af . heterostipum fi;om Brazil. 



\T}\ ) UO^A, 




M. Fulford _162 Bazzania 

basis on which to establish a separate species. Although the type 
plants of M . Richardianum collected in the Straits of Magellan had 
underleaves with the hyaline cells larger and more or less irregu- 
larly arranged, the other South American material showed a whole 
series of variation in size and the arrangement of these cells. 

Plants collected by ULB in Brazil, in the STEPHANI Herbarium 
at Harvard University under the name of M. heterostipum (FiG. 
58, nos, 15-17), have been taken as the type of that species. 
STEPHANI (1909, 532) gives as the habitat "America tropica et 
subtropica, haud rara", and does not mention a collector's name. 
The underleaves of most of these plants have the teeth well devel- 
oped, the cells mostly of the smaller size and with thickened walls. 
The other variation can also be found. 

DISTRIBUTION: Brazil: without locality, Ule, the type (?) of Af. 
heterostipum (H) ; Organ Mountains, Gardner (NY). Peru: without 
locality, "Hb. Dr. Gottsche", a part of the material referred to by Lindenberg 
(G. L. & N. p. 215), (NY, H) ; Tatanara, Lechler (NY); without locality, 
Jameson (NY). Chile : ChiloS Island, Captain King (NY). Pata- 
gonia: Albert Bay, Dr. Coppinger (NY); Straits of Magellan, Richard, 
the type of M. Richardianum (NY). Also Central Africa, South Africa and 
Australia. The original material of J. convexa was collected at the Cape of 
Good Hope by C. P. THUNBERG. 

REFERENCES: Hooker (1867, 624); Lehmann (1829, 364); Mitten (1855, 
147); Schwaegrichen (1814, 22); Sprengel (1827, 225); Stephani (1909, 532, 
534; 1 cones, Mastigobryum nos. 439, 443, 444). 

46. Bazzania Stephani (Jack) comb. nov. 

Mastigobryum Stephani Jack, in Stephani, Hedw. 25: 235. pi. 1, fig. 10-12. 
1886. 

Plants scattered among mosses, pale yellow-green to brownish : 
stems very slender, to 3 cm. in length, with leaves to 1.5 mm. 
broad, prostrate; the lateral branches occasional, diverging $t a 
wide angle, ventral branches frequent, usually leafy: rhizoids oc- 
casional from the bases of the underleaves of ventral branches: 
the line of leaf insertion little curved in its upper part; the leaves 
vittate, approximate to subimbricated, spreading, plane* the teeth 
and a little of the apical portion often deflexed, unsymmetrically 
oblong-ovate, 0.7 mm. - 0.9 mm. long, mostly 0.35 mm. broad at the 
base, narrowed a little to the transversely truncate, obscurely tri- 
dentate apex; the dorsal margin convex from a straight base, 
covering one-third the stem, the ventral margin nearly straight, 
the base scarcely dilated, the apex obscurely three-toothed or -lobed, 
the teeth one or two cells high, two to four cells broad at the base, 
the sinuses lunulate, the margins entire; the leaf cells quadrate to 
rectangular in outline, with uniformly thickened walls, the cell 
lumina rounded, the vitta of three or four rows of enlarged cells 
clearly differentiated, the cuticle verruculose; cells of the apical 
portions and dorsal base averaging 16/x X 16/x, those of the vitta 
to 32/i X 24/A, the walls thin, the trigones conspicuous : the under- 
leaves large, approximate to imbricated, hyaline above, attached in 
a straight line, rectangular in outline, averaging 0.35 nun. - 0.42 




FIG. 59. Bazzania Stephani (Jack) Fulford. 1. Portion of plant, 
ventral view, X 30. 2. Underleaves. NOB. 1-2 drawn from plants of Burchell, 
Cat. Geog. PL Brasil. Trop. No. 8847. 



M. Fulford 164 Bazzania 

mm. long X 0.28 mm. wide, the apex undulate, two- to four-lobed 
or -toothed, the lateral margins entire, the chlorophyllose cells 
restricted to a small, interior, basal area, the cells all quadrate to 
rectangular in outline, the walls uniformly thickened: sexual 
branches not seen. 

HABITAT: Over rocks and soil with other bryophytes. 

The distinguishing characteristics of the species are its small 
size, the yellow-green color, the plane, obscurely tridentate, vittate 
leaves, and the large, elongate underleaves, often two- or four- 
lobed or -toothed at the apex, and hyaline to near the base. The 
cuticle is verruculose. (FiG. 59, nos. 1-2). 

The plants are very similar to B. convexa. The leaves of the 
two species are identical. However, the underleaves of B. convexa 
are medium size and to one-half divided into four slender teeth, 
while those of B. Stephani are very large and divided to one-fifth 
or less into two or four lobes which are often obscure. Both species 
have the chlorophyllose cells of the underleaves restricted to a 
small basal area. 

DISTRIBUTION : Brazil: without locality, Burchell, Cat. Geog. PL Brasil. 
Trop. 3847 (NY). The original material was collected on Johanna Island of 
the Comoro group in the Indian Ocean. 

REFERENCES: Stephani (1909, 535; Iccmes, Mastigobryum no. 441). 



EXCLUDED SPECIES 

Mastigobryum mutants Herzog, n. sp. Repert. Spec. Novarum Regni Veg. 
21: 26. pi. XI, fig. 2, 1925, Brazil, belongs to some other genus (Isotachis ?). 

SPECIES NOT AVAILABLE FOR THIS STUDY 

Because of the unsettled world conditions and the war, it has 
been impossible to borrow some of the type material of South 
American species from the European Herbaria. A study of these 
species and varieties will no doubt reveal many synonyms among 
them. The species and varieties not included are as follows :- 

Bazzania adnexa (L. et L.) Trevis. reported from Chile by HERZOG. Arch. Esc. 

Farm. Fac. Cien. Med. C6rdoba 1938. No. 7. p. 26. 1938. 
Mastigobryum arcuatum Lindenberg & Gottsche, in G. L. & N., Syn. Hep. 718. 

1847. Mexico. 

Mastigobryum azuayense Stephani, Spec. Hep. 6: 454. 1924. Ecuador. 
Bazzania bidens var. heterodonta Spruce, Trans. & Proc. Bot. Soc. [Edinburgh] 

15: 372. 1885. Peru. 

Mastigobryum bogotense Stephani, Hedwigia 24: 246. 1885. Colombia, 
Mastigobryum brasiliense Gottsche & Linden-berg, in G. L. & N., Syn. Hep. 227. 

1845. Brazil. 
Mastigobryum Breutelianum var. guadeloupense Bescherelle, Jour. Bot. 7: 187. 

1893. Guadeloupe, (nomen nudum). 

Mastigobryum Burchellii Stephani, Spec. Hep. 3: 509. 1908. Magellan Straits. 
Mastigobryum cerinum Stephani, Spec. Hep. 3: 457. 1908. 
Mastigobryum columbicum Stephani, Spec. Hep. 6: 458. 1924. Colombia. 
Mastigobryum creberrimum Stephani, K. Svenska Vetensk. Handl. 46: no. 9. 

60. 1911. 

Mastigobryum decurrens Stephani, Bibliotheca Bot. 87*: 223. 1916. Bolivia. 
Mastigobryum ecuadorense Stephani, Spec. Hep. 6: 461. 1924. 
Mastigobryum heterophyllum Stephani, Spec. Hep. 6: 466. 1924. Ecuador. 
Bazzania hum if usa var. olivacea Spruce, Trans. & Proc. Bot. Soc. [Edinburgh] 

15: 380. 1885. Peru. 
Mastigobryum portoricense var. laxa Bescherelie, Jour. Bot. 7: 186. 1893. 

Guadeloupe, (nomen nudum). 
Mastigobryum Schlimianum Gottsche, Ann. Sci. Nat. V. Bot. 1: 140. 1864. 

Colombia. (The portion* of the type in the Farlow Herbarium is too frag- 
mentary for determination.) 

Herpetium scutigerum Montague, Ann. Sci. Nat. II. Bot. 9: 44. 1838. Peru. 
Mastigobryum venezuelanum, Molkb. ms. in Sande Lacoste, Syn. Hep. Javan. 

104. 1857. Venezuela. 
Mastigobryum venezuelanum Stephani, Spec. Hep. 3: 530. 1909. Venezuela. 

(Fendler.) 

Bazzania vincentina var. macrophylla Spruce, Trans. & Proc. Bot. Soc. [Edin- 
burgh] 15: 378. 1885. Ecuador. 



DISTRIBUTION 

Few accurate statements concerning the distribution of the 
species of Bazzania can be made at this time, since our present 
knowledge of the occurrence of the plants represents a "distribu- 
tion of collectors", and unfortunately few adequate collections have 
been made in most areas. 

The Bidentatae group seems to be restricted to the West Indies 
and northern South America. Bazzania Herminieri and perhaps 
B. platystipula may be endemic to the West Indies, and B. rorai- 
mensis seems to be limited to Mt. Roraima in British Guiana. The 
other species are rather widely distributed within the area, with 
B. gracilis most abundant. 

Most of the members of the Grandistipulae Section occur 
throughout the West Indies, often in Mexico and Central America, 
and are abundant in tropical South America. Among the most 
frequently collected species may be mentioned B. stolonifera, B. 
Breuteliana, B. jamaicensis and B. longistipula. At the present 
time B. pallide-virens, B. Glaziovii and B. tricuspidata are known 
only from restricted areas in South America and may be localized 
endemics. B. longa is very abundant throughout the West Indies 
but no specimens from Mexico, Central, or South America have 
come to light. 

The Connatae Section contains three distinct geographic segre- 
gates; the one, made up of B. Schwaneckiana, B. pycnophylla, B. 
Eggersiana, B. armatistipula and B. cubensis, in which the leaf 
apices are serrate and the underleaf is connate with one leaf by 
only a few cells, is restricted to the West Indies, with some of the 
species further limited in their distribution to one or two of the 
islands ; another, B. Fendleri, in which the underleaf is also connate 
with only one leaf but differing from the above group in habit, 
color, cells, etc., occurs in Brazil, Ecuador, and Venezuela; and the 
third, including B. peruviana, B. Skottsbergii and related species, 
in which the underleaf is connate with a pair of leaves, occurs south 
of the equator from Peru south to the Straits of Magellan. The 
species of this latter group show close relationships with species 
from South Africa and New Zealand, and in some instances may 
even be identical. 

The Appendiculatae Section is made up of a group of closely 
related species which are most abundant in tropical and subtropical 
South America. B. teretiuscula is also abundant in Central 
America and Mexico; B. asperistipula seems to be restricted to 
northern South America and Central America ; B. falcata and B. 
Hookeri in addition to a widespread distribution in South America 
occur also in the West Indies ; B. robicsta and B. acanthostipa are 



Bazzania 



167 



Distribution 



fairly abundant but restricted to South America; and B. canelensis 
seems to be an endemic of the mountains of Ecuador. 

Three of the four members of the Vittatae Section are known 
to occur also in Africa, New Zealand or Australia. B. Tayloriana 
occurs in Colombia and New Zealand; B. Stephani in Brazil and 
Johanna Island of the Comoro group off the coast of Africa ; and 
B. convexa, from Brazil, Peru, Chile and Patagonia, is also abun- 
dant in central and southern Africa, New Zealand and Australia. 
B. Spruceana seems to occur only in Peru. 




LITERATURE 

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Antilles franchises (Guadeloupe et Martinique). Journal de Botanique 

7: 174-180; 183-194. 
BOSWELL, HENRY. 1887. Jamaica Mosses and Hepaticae. Journal of Botany 

25: 45-50. 
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Commentationes Biol. 3: 5. 1928. 
. 1929. Eine neue moossystematische Methodik nebst einigen ihrer 

Resultate und ein neues Nomenklatursystem. Skand. Naturforskermode 

18: 225-229. 
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15 pi. 
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1-24. Tournay. 

Duss, R. P. 1903, 1904. See STEPHANI, F. 1903, 1904. 
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Bazzania 169 Literature 

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M. Fulford 170 Bazzania 

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Icones Hepaticarum. Mastigobryum. Nos. 1-465. Unpublished 



drawings made by FR. STEPHANI; recently several sets of tracings of these 

have been made and distributed by Miss JOHANNA STEPHANI of Leipzig. 
SWARTZ, O. 1806. Flora Indiae occidentalis. Hepaticae. 3: 1842-1885. 

Erlangen. 
TAYLOR, TH. 1846. New Hepaticae. London Journal of Botany 5: 258-284; 

365-417. 
WALLROTH, C. F. G. 1831. Flora Cryptogamica Germaniae. Pars Prior, i- 

xxvi, 1-654. Numberg. 

WEBER, FR. 1815. Historiae Muscorum hepaticorum prodromus. 1-160. Kiel. 
WEBER, FR. und MOHR, D. M. H. 1807. Handbuch der Einleitung in das 

studium der kryptogamischen Gewachse. i-xlvi, 1-509. 12 pi. Kiel. 



INDEX 



Synonyms are listed ito italics, descriptions of the species in bold face type, 
and figures m italics. 

Acromastigum 3 

AppendiciUata 1, 3, 130 

Appendiculatae 4, 7, 37, 38, 125, 132, 137, 152, 154, 166 

Bazzania 1, 4, 5, 6, 7, 122, 166 

acanthostipa 125, 148, 149, 151, 166 

acuminata 38, 39, 74, 75, 76, 86 

adnexa 165 

affinis 38, 39, 41, 42, 50 

ambigua 8, 22 

ancistrodes 129, 132, 133, 134, 135 

armatistipula 105, 108, 114, 115, 166 

asperistipula 125, 126, 128, 134, 135, 166 

aurescens 38, 63, 64, 67 

bidens Trevis. 8, 9, 10, 14, 18, 24, 35 

bidens Spruce 11, 30, 33, 35, 36 

bidens var. 30, 36 

var. dissodonta 11, 14, 15 

var. heterodonta 165 
var. vittata 30, 35 



Breuteliana 38, 50, 54, 68, 69, 71, 72, 77, 86, 90, 166 

canelensis 37, 125, 152, 153, 167 

chilensis 38, 51, 53 

chimborazensis 68, 70, 71, 73, 74, 75, 77 

conchophylla 144, 145, 150 

consanguinea 81 

convexa 155, 159, 161, 164, 167 

cubensis 105, 108, 112, 116, 117, 166 

cuneistipula 8, 9, 18, 19, 21, 23, 29 

decidua 81, 84, 85, 86, 87 

denticulata 38, 56, 57, 59 

denudata 8 

dissodonta 11, 12, 13, 15 

diversicuspis 37, 38, 77, 78, 81 

Eggersiana 105, 108, 110, 113, 116, 166 

falcata 125, 129, 131, 133, 139, 142, 166 

Fendleri 103, 104, 105, 166 

flavicans 135, 138, 139, 140 

Glaziovii 38, 65, 66, 166 

Gottscheana 90 

gracilis 8, 9, 11, 15, 30, 31, S3, 166 

Herminieri 8, 9, 11, 15, 17, 24, 166 

heteroclada 144, 146, 147, 148, 150 

Hookeri 38, 125, 135, 1 36, 138, 142, 166 

Hookeri var. 63, 64, 65 

humifusa 144, 145, 150 

olivacea 165 



jamaicensis 39, 96, 97, 99, 102, 166 

var. chamaecardia 96, 98, 100 

Krugiana 81, 84, 85, 86, 87 
latidens 39, 88, 89 
leptostipa 36, 44, 46, 47, 48, 51 
Liebmanniana 125, 142, 143 



M. Fulf ord 172 Bazzania 

- Lindigii 44 

- longa 37, 39, 90, 91, 9S, 95, 166 
-- var. papillata 94, 95 

- longistipula 11, 24, 37, 39, 81, S3, 87, 90, 160, 166 

- longistipula vwr. 81 

var. polymastix 81, 84, 86 



loricata 22 
novae-zelandia 123 
otites US, 150 

pallide-virens 38, 42, 43, 166 

peruviana 103, 119, 1*0 y 122, 123, 166 
phyllobola 8, 11, IS, 18, 24, 36, 81 
planiuscula 56 

platystipula 8, 25, 26, 29, 166 

portoricenaia var. pycnodictyon 68, 70, 71, 74 
pycnophylla 105, 108, 109, 110, 111, 112, 114, 116, 166 
quadricrenata 38, 60, 61, 65, 67 

robusta 38, 125, 140, 141, 166 

roraimensis 8, 27, 28, 166 

Rusbyi 56, 58, 59, 60 

Schwaneckiana 37, 105, 106, 107, 110, 112, 118, 166 

Skottsbergii 103, 122, 166 

apinigera 144, 146, 147, 148, 150 

Spruceana 155, 156, 167 

Stephani 155, 162, 163, 167 

stolonifera 38, 44, 45, 47, 49, 52, 166 

subfalcata 90 

superb a 135 

taleana 38, 39, 54, 55 

Tayloriana 155, 157, 158, 167 

tenera 19 

teretiuscula Trevis, 125, 144, 145, 147, 166 

teretiuscula Spruce 81, 82, 8S, 84, 85, 86, 150 

trichodea 30, S3, 35, 36 

tricrenata 8, 84 

tricuspidata 39, 79, 80, 86, 166 

trilobata 1, 46 

vtincentina 44 

- f. defolians 51 

- f. minor 51 

- f. ramulosa 51 

- f. rufeacena 51 

- var. macrophylla 165 

- var. aubed&ntata 152, 153, 154 

- var. aubmutica 152, 154 

var. subrectifolia 68, 75, 135, 138, 139, 140 



viridiaaima 68, 70, 71, 73, 74, 75 
Wrightii 39, 51, 74, 98, 100, 101 



Bidentata 1, 2 

Bidentatae 4, 7, 8, 27, 29, 32, 166 

Classification 4, 7, 37 

Connata 1, 3, 37 

Connatae 4, 7, 37, 60, 103, 108, 109, 116, 118, 166 

Cordiatipula 1, 3, 7, 37, 130, 139 

Distribution 166 

Fiaaistiptda 1, 3, 110, 119 

Fissistipulae 4, 37, 124 

Frullania 109 

Grandiatipida 1, 2, 3, 37, 114 

a eonnata 1, 2 

ft libera 1, 2 



Bazzania 173 Index 

Grandistipulae 4, 7, 37, 38, 56, 114, 124, 166 
Herpetium I, 5 

sect. Mastigobryum 5 

jamaicfinse 96, 98 

scutigerum 365 

stolaniferum 44 

var. bidens Mont. 35 

var. bidens Nees 9 

var. vrreguLare 51 



vincentianum 44, 51 



Inaequilatera 1, 3 
Introduction 1 
Integrifolia 1, 2, 3, 7, 108 
Jungermannia 1 

cvnvexa 159, 160, 161, 162 

longa 90, 92, 94 

nitida 159 

peruviama 119, I0 

stolonifera Sieb. 90 

stolanifera Swartz 44, 45, 48, 90 

tridens 9 

trilobata 1 

vincentiama 44, 46, 47, 48, 51 

Keys 7, 8, 37, 38, 103, 124, 125, 155 
Literature 168 

Lophozia 6 
Mastigobryum 1, 5 
acantkostipum 151 

acuminatum 75, 77 

affine L. & G. 39, 40 

affme Mitten 39 

andstrodes 129 

arcuatum 94, 165 

armatistipulum 114, 116 

armatum 129, 132, 133, 134, 135 

asperistipulum 125 

aurescens 63 

azuayense 165 

bidens 9, 30, 35 

bogotense 165 

bolivianum 140, 142 

brachyphyllum 81, 84 

brasiliense 94, 165 

Braitniamum 135, 138, 139 

Breuteliamum 68, 73, 74 

var. guadeloupense 165 
brevifolium 19, 22, tS, 24 
Burchellii 124, 165 
eanelenae 152, 154 
caracanum 96, 98, PP. 100 
cerinum 122, 123, 165 
chamaecardion 96, PP 
okilense 51 
chimborazense 68 
columbicum 165 
consanguineum 81, 84, 85, 86 

var. brachyphyUum 81, 85, 86 

convexum 159 
oortioela 19, 22, t3, 24 
creberrimum 165 
cubense 116, 119 



M. Fulf ord 174 Bazzania 



Cuervi 68, 70, 71, 73, 74 

cunei folium 30, 35, 36 

cuneistipulum 19, 22, 24 

deciduum 37, 81 

decurrens 165 

denticuLatum 56, 58, 60 

disaodontum 11, 15 

diversicuspe 7, 77 

Doumt 135, 139, 140 

ecuadorense 165 

Eggersianum 110 

elegantulum De Not. 81 

elegantulum Gottsche 24, 37, 81, 84, 85, 86, 87 

falcatum 129, 134 

flavioans 135 

Fendleri 105 

Glaziovii 65 

Gottackeanum 90, 92, PJ, 94, 96 

gracile 30 

trichodea 32, 35 

guadaloupense 135, 13#, 139 
Hansenii 81, 84, 85, 86, 7 
Hwriotii 68, 73, 74 
Herminieri 15 

f. applanata 18 

f . /attar 18 

var. brevifolia 18 



heterockidum 144 

heterophyllum 124, 165 

heterostipum 159, 160, l^J, 162 

Hookeri 135, 139 

humifuswn 144 

inciso-bilobatum 39, 40, 42 

indaostijndum 144, 150 

jamaicense 96 

Krugicmum 37, 81 

latidems 88 

Lechleri 119, J20, 122 

leptostipum 44 

Liebmamnianum 142 

Lvndigii 44, 46, -47, 50, 51 

longicuspe 54, 55, 56 

longistipulum 81, 84, 85, 86 

kmgum 90 

Mwtianum 60, 01, 62, 74 

Miillerianum 75, 70, 77 

mutans 165 

novae-zelandiae 123 

orizabense 75, 70, 77 

pallide-vi/rens 42 

paludosum 81, 82, 5, 84, 86 

papillatum 94, 05 

30, W, 35, 86 

119 
var. mmwum 119, 122 

phyllobolum Gottsche 81, 84 

phyllobolum Steph. 11, 36 

planiusculum 56, 58, 50, 60 

portoricense 50, 68, 70, 71, 73, 74, 139 

var. laxa 165 



Puiggwm 81, 84, 85, 86, 57, 88 



Bazzania 175 Index 



pycn&phyUum 109 
qitadricrenatwrn 60 

f. pauper euLa 60, 61, 62 



Quelchii 44, 46, 47, 50, 51 

Rwhardumum 159, 160, 161, 162 

robustum 140 

roraimense 27 

saxatile 81, 82, , 84, 85, 86 

Schlimianum 140, 165 

Schwanechianum 2, 3, 7, 106 

scutigerum 65 

Skottsbergii 122 

spectosum 90, 94, 95, 96 

spmigerum 144 

Spruceanum 155 

Stephcmi 162 

stoloniferum 44 

/? intermedia 51 

var. cubense 100, JTW, 102 

var. grcmatensis 51 
subfalcatum 90, 94, P5, 96 
superbum 135, j(J0, 139 
sylvaticum Gottsche 44, 68, 74 
sylvaticum Steph. 44, 46, 47, 50, 68 
taleanum 54, 55, 56, 74 
Taylorianum 157 

18, 19, 22, 25, 24 
terme 37, 90, 0S, 94 
teretiusculum 81, 129, 144, 148, 150 
tocuticmum 129, 132, .m, 134, 135 
trichoideum 30, 35 
tricuspidatum 2, 37, 79 
tritnitatis 90, 94, P5, 96 
Tilrkheimii 44, 46, 47, 50, 51 
Uleanum 68, 70, 7jf, 74 
vorta6t/e 18, 19, 22, 23, 24 
vartedentatum 54, 56 
venezuelanum Molkb. 165 
venezuelanum Steph. 165 
verrucosum 135, 138, 139, 140 
vincentinum 44 
viridissimum 51, 68 
Wrightii 100, 102 



Pwrvistiptda 1, 2, 7, 37 
Pleuroschisma 1, 5 

stoloniferum 44 

Scapania 6, 50 
Serrulate, 1, 3, 37, 116 

a connata 1, 3 

Ji&era 1, 3 

Subgenera 7, 8, 37 
Tridentata 1, 2 
Tridentatae 4, 7, 8, 24, 37, 60 
Vittata 1, 3 

Vittatae 4, 7, 37, 155, 167